Short-Term Response of Fleshy Fungi to Prescribed Burning in a Minnesota

Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations

1-1-2002

Short-term response of fleshy fungi ot prescribed burning in a Minnesota near-boreal/Great Lakes pine forest

Erin Jane Heep Iowa State University

Follow this and additional works at: https://lib.dr.iastate.edu/rtd

Recommended Citation Heep, Erin Jane, "Short-term response of fleshy fungi ot prescribed burning in a Minnesota near-boreal/ Great Lakes pine forest" (2002). Retrospective Theses and Dissertations. 19871. https://lib.dr.iastate.edu/rtd/19871

This Thesis is brought to you for free and open access by the Iowa State University Capstones, Theses and Dissertations at Iowa State University Digital Repository. It has been accepted for inclusion in Retrospective Theses and Dissertations by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. Short-term response of fleshy fungi to prescribed burning

in a Minnesota near-boreal/Great Lakes pine forest

Erin Jane Heep

A thesis submitted to the graduate faculty

in partial fulfillment of the requirements for the degree of

MASTER OF SCIENCE

Major: Botany (Mycology)

Program of Study Committee: Lois H. Tiffany (Major Professor) Thomas W. Jurik J. Michael Kelly

Iowa State University

Ames, Iowa

2002 11

Graduate College Iowa State University

This is to certify that the master's thesis of

Erin Jane Heep has met the thesis requirements of Iowa State University

Signatures have been redacted for privacy 111

For Tom

Pine mushroom - some kind of leaf sticking to it. Matsuo Basho ( 1644-1694) IV

TABLE OF CONTENTS

CHAPTER 1. GENERAL INTRODUCTION 1 Introduction 1 Thesis organization 2 References 3

CHAPTER 2. EPIGEOUS BASIDIOCARP SURVEY FOLLOWING 5 PRESCRIBED BURNING IN A NEAR-BOREAL/GREAT LAKES PINE FOREST IN MINNESOTA Abstract 5 Introduction 6 Study site and methods 10 Results and discussion 14 Acknowledgments 3 3 References 3 4

CHAPTER 3. SHORT-TERM EFFECTS OF PRESCRIBED BURNING ON 42 THE ECTOMYCORRHIZAL FUNGAL COMMUNITY IN A NEAR-BOREAL/GREAT LAKES PINE FOREST IN MINNESOTA Abstract 42 Introduction 4 3 Study site and methods 46 Results and discussion 50 Acknowledgments 5 5 References 5 6

CHAPTER 4. GENERAL CONCLUSIONS 60 General discussion 60 Recommendations for future research 63 References 64

APPENDIX A. IDENTIFICATION RESOURCES 66

APPENDIX B. ADDITIONAL TABLES 72

APPENDIX C. MAPS 96 V

ACKNOWLEDGMENTS

I could not have completed this thesis without the help of many people along the way. I would like to extend my warmest thanks to the following people: Dr. Lois H. Tiffany for offering support and encouragement to me when I felt overwhelmed, for sharing a portion of her vast knowledge, for her suggestions, advice, time, and for always cheerfully and willingly answering my many, questions. Dr. J. Michael Kelly for his willingness to read endless revisions, for helping me to maintain perspective focus, and for always challenging me to "think about the take-home message". Dr. Thomas W. Jurik for reading rough drafts and offering thoughtful and very helpful suggestions. Dr. David McLaughlin at the University of Minnesota for all his suggestions, and for answering many email questions promptly and thoroughly. Patrick R. Leacock for writing a fabulous dissertation on the diversity of northern Minnesota's ectomycorrhizal fungi - it was largely that dissertation that inspired me to pick the topic of my own research. I also appreciate the information he provided on the Lactarius species that were documented during the study. I would also like to extend very special thanks to the many USDA Forest Service scientists of the Superior National Forest who have supported me from the very beginning, especially Robert Kari (soils), Jim Hinds (fire), and Don Potter (biology). They answered many questions throughout, challenged me with questions in return, made many resources available to me, and willingly shared their technical expertise. Sincere thanks to Dan Hanson of the Minnesota Department of Natural Resources for his valuable help in describing the soil, landscape, and geological characteristics of the study site. Thank you to my parents, Bert and Diane, for their tremendous support, and for periodically assisting me with the fungal surveys. I am forever grateful to Tom for accompanying me on nearly all the collecting trips, for checking Unit 3 the summer after my knee surgery, for his willingness to listen, and for stimulating, thought-provoking discussions on fire ecology, ectomycorrhizal fungi, and the coniferous forest ecosystem. Thank you for patiently waiting for me to join you in the northwoods upon the completion of this thesis. 1

CHAPTER!. GENERALINTRODUCTION

Introduction

Fungi play significant roles in the ecosystem through nutrient cycling, food webs, and forest diseases and through ectomycorrhizal mutualisms that influence seedling survival, tree growth, and overall forest health by enhancing nutrient acquisition, drought tolerance, and pathogen resistance of their hosts (Molina et al. 2001; Peter et al. 2001). Despite their importance, fungi are not usually mentioned in discussions of biodiversity and conservation

(Hawksworth 1992). Yet, declines in fungal diversity, primarily the result of human-induced disturbances, have been documented in Europe since the 1980s (Arnolds and de Vries 1993).

It is difficult to determine whether declines in species diversity or range changes are occurring in North America as they are in Europe due to the lack of baseline data and accurate distribution maps for North America (Redhead 1989; Jaenike 1991).

Fire, both natural and prescribed, is one of the main disturbance events in the coniferous forests of northeastern Minnesota. Prescribed burning, a management tool used by land managers to mimic low intensity natural fires, has increased substantially in the past two decades (Neary et al. 1999). While several studies have investigated the effects of wild or prescribed fire on fungal communities in other countries or ecosystems (e.g.,O'Halloran et al. 1987; Torres and Honrubia 1997; Stendell et al. 1999; Dahlberg et al. 2001), no one has attempted to determine the effects of prescribed fire on the fungal communities in northeastern Minnesota. Due to variation amongst ecosystems caused by fuel types, species adaptation to fire, and abiotic factors, it is difficult to estimate the ecological impacts of fire 2 on one ecosystem and apply them to another (Herr et al. 1994). Therefore, to understand the biological and functional diversity of forest fungi, and before land managers in Minnesota can incorporate fungi into their management and conservation plans, fundamental information on the fungal species within the forest community and an understanding of the impacts of natural and anthropogenic disturbance must first be obtained.

A comprehensive picture of the fungal communities in northeastern Minnesota is needed as a basis for detecting changes in those communities. One goal of this study was to obtain quantitative baseline data on species abundance, species richness, and fruiting phenology of the epigeous macrofungal community in a near-boreal/Great Lakes pine forest located in the Superior National Forest. Another goal of this study was to identify potential species composition differences in the fungal community following prescribed fire. The final goal was to evaluate, at the same location, the short-term effects of prescribed burning on the ectomycorrhizal fungi by comparing above-ground ectomycorrhizal species richness and sporocarp abundance in burned and unburned study units.

Thesis organization

In addition to the general introduction and conclusion, this thesis is composed of two main chapters, each of which is an individual manuscript that has been submitted to a professional journal. Chapter 2 summarizes the baseline data of the three-year study, for topics such as species richness, sporocarp abundance, sporocarp density, and phenology. The effects of fire on the fungal community composition are also investigated. Chapter 3 emphasizes the effects that prescribed burning had on ectomycorrhizal fungi in particular.

Differences between burned and unburned units, in terms of overall sporocarp abundance, 3 ectomycorrhizal sporocarp abundance, ectomycorrhizal species richness, and the percentage of ectomycorrhizal species, are all discussed in this chapter.

Three appendices are also included at the end of the thesis. Appendix A lists several resources used in the identification of the fleshy, epigeous basidiocarps collected for this study. Copies of data forms used for noting macro- and microscopic characters are also included in Appendix A. Appendix B presents various combinations of the data collected during this study in the form of additional tables that are not included in Chapters 2 or 3.

Appendix C consists of detailed maps of the study site and the burned study units. The maps were created by the Fire Management Officer at the Superior National Forest, USDA Forest

Service.

References

Arnolds, E., and de Vries, B. 1993. Conservation of fungi in Europe. In Fungi of Europe: investigation, recording, and conservation. Edited by D.N. Pegler, L. Boddy, B. Ing, and P.M. Kirk. Royal Botanic Gardens, Kew. pp. 211-230.

Dahlberg, A., Schimmel, J., Taylor, A.F.S., Johannesson, H. 2001. Post-fire legacy of ectomycorrhizal fungal communities in the Swedish boreal forest in relation to fire severity and logging intensity. Biol. Conserv. 100: 151-161.

Hawksworth, D.L. 1992. Fungi: a neglected component of biodiversity crucial to ecosystem function and maintenance. Canadian Biodiversity, 1: 4-10.

Herr, D.G., Duchesne, L.C., Tellier, R., McAlpine, R.S., and Peterson, R.L. 1994. Effect of prescribed burning on the ectomycorrhizal infectivity of a forest soil. Int. J. Wildland Fire, 4: 95-102.

Jaenike, J. 1991. Mass extinction of European fungi. Trends Ecol. Evol. 6: 17 4-17 5. 4

Molina, R., Pilz, D., Smith, J., Dunham, S., Dreisbach, T., O'Dell, T., and Castellano, M. 2001. Conservation and management of forest fungi in the Pacific Northwestern United States: an integrated ecosystem approach. In Fungal conservation: issues and solutions. Edited by D. Moore, M.M. Nauta, S.E. Evans, and M. Rotheroe. Cambridge University Press, Cambridge. pp. 19-63.

Neary, D.G., Klopatek, C.C., DeBano, L.F., and Ffolliott, P.F. 1999. Fire effects on belowground sustainability: a review and synthesis. For. Ecol. Manag. 122: 51-71.

O'Halloran, K.A., Blair, R.M., Alcaniz, R., and Hershel, F.M., Jr. 1987. Prescribed burning effects on production and nutrient composition of fleshy fungi. J. Wildl. Manag. 51: 258- 262.

Peter, M., Ayer, F., Egli, S., and Honegger, R. 2001. Above- and below-ground community structure of ectomycorrhizal fungi in three Norway spruce (Picea abies) stands in Switzerland. Can. J. Bot. 79: 1134-1151.

Redhead, S.A. 1989. A biogeographical overview of the Canadian mushroom flora. Can. J. Bot. 67: 3003-3062.

Stendell, E.R., Horton, T.R., and Bruns, T.D. 1999. Early effects of prescribed fire on the structure of the ectomycorrhizal fungus community in a Sierra Nevada ponderosa pine forest. Mycol. Res. 103: 1353-1359.

Torres, P., and Honrubia, M. 1997. Changes and effects of a natural fire on ectomycorrhizal inoculum potential of soil in a Pinus halepensis forest. For. Ecol. Manag. 96: 189-196. 5

CHAPTER 2. EPIGEOUS BASIDIOCARP SURVEY FOLLOWING PRESCRIBED BURNING IN A NEAR-BOREAL/GREAT LAKES PINE FOREST IN MINNESOTA.

A paper submitted to the Canadian Journal of Botany

Erin J. Heep

Abstract

I collected quantitative baseline data on the soil-borne, epigeous basidiocarp community in a near-boreal/Great Lakes pine forest located in the Superior National Forest,

Minnesota. Prescribed burned and unburned units were surveyed in 1999, 2000, and 2001, and were compared in terms of sporocarp abundance, overall fungal species richness, ectomycorrhizal species richness, percent ectomycorrhizal species, and fruiting phenology.

A total of 179 taxa was documented, representing 33 genera and four orders (Agaricales,

Boletales, Cortinariales, and Russulales). Ninety-three percent were true mushrooms and 7% were boletes. There were more ectomycorrhizal species present than non-mycorrhizal species. The dominant genera were Russula, Cortinarius, Lactarius, and Amanita. The burned units exhibited less species richness and less ectomycorrhizal species richness than the unburned units. However, the percentage of ectomycorrhizal species in burned units did not differ from the unburned units. There was no difference between the burned and unburned units in terms of sporocarp abundance. Sporocarp fruiting was most productive in

September. In general, the low intensity burns did not appear to have a long-term adverse effect on the fungal community in the near-boreal/Great Lakes pine forest in northeastern

Minnesota.

Key words: fire, fungi, ectomycorrhizal, saprobic, carbonicolous 6

Introduction

Fungi are rarely mentioned in discussions of biodiversity and conservation

(Hawksworth 1992). However, there is a growing effort by mycologists to educate scientific and lay communities alike on the benefits of and need for global fungal conservation (Moore et al. 2001). Europe has been in the forefront of fungal conservation since the detection of declines in macrofungal populations and shifting ranges during the past few decades

(Arnolds 1991; Ing 1996). The concept that fungi may be threatened and deserve conservation attention equal to that placed on animals and vascular plants is now well established in Europe (Courtecuisse 2001), but less so in North America.

Fungal declines have been, and continue to be, caused mainly by destruction of appropriate habitat. However, pollution, climate change, habitat fragmentation, and changes in land use are also recognized as causes of decline (Ing 1996; Courtecuisse 2001).

Controversy still surrounds the issue of wild mushroom harvesting, and potential correlations between long-term harvesting and species decline have not been thoroughly investigated

(Rotheroe 1998; Arora 2001; Pilz and Molina 2002).

While fungal conservationists have little or no control over global air pollution or habitat fragmentation, Courtecuisse (2001) recommends that mycologists raise awareness of the value of fungi at the ecosystem level by using "tools" such as inventories and checklists, mapping programs, and ultimately, Rarity, Endangerment, and Distribution Data (Red Data) lists. European Red Data lists have been compiled for the continent (Ing 1993a, 1993b) and many countries, including, but not limited to, Great Britain (Ing 1992), France (Courtecuisse

1997), Hungary (Siller and Vasas 1995), The Netherlands (Arnolds 1997), Norway 7

(Bendiksen and H0iland 1992), Poland (Wojewoda and Lawrinowicz 1992), Slovakia (Lizon

1995), and Switzerland (Senn et al. 1997).

It is estimated that there are at least 1.5 million fungal species worldwide

(Hawksworth 2001 ). Yet, so few species have been described that many may be threatened before their existence is known (Hawksworth 1992; Staley 1997). In North America, vast areas remain unexplored mycologically, and accurate distribution maps are often not available (Redhead 1989; Jaenike 1991). Our knowledge of fungal diversity is so limited, we do not know if and when most species are threatened (Staley 1997). Without baseline data, it is difficult to determine whether species diversity declines or range changes are occurring in

North America as they are in Europe (Jaenike 1991). The Pacific Northwest region of the

United States is one area of North America that has been relatively thoroughly explored, and preliminary Red Data lists have been compiled for Oregon (Castellano 1997 a, 1997 b ), Idaho

(Castellano 1998), and Washington (Washington Natural Heritage Program 1997).

The near-boreal forest of Minnesota is an area of North America that has been quite neglected mycologically. The diversity and distribution of Minnesota macrofungi are largely unknown, and only the central and eastern parts of the state have been studied extensively

(McLaughlin and Leacock 1994). Approximately 1,000 species ofmacrofungi have been recorded for Minnesota, yet it is estimated that the number of species actually present is more than 3,000 (McLaughlin and Leacock 1994). There are few published studies documenting fungal communities in northern Minnesota. Weaver and Shaffer (1969, 1972) compiled lists of Minnesota fungi collected in various areas of the state including northern locales in Itasca

State Park, Itasca County, and St. Louis County. Doudrick et al. (1990) surveyed black spruce stands in northern Minnesota for ectomycorrhizal fungi and documented 46 species. 8

Leacock ( 1997) surveyed ectomycorrhizal fungi in old growth and young red pine and northern hardwood-conifer forests in Scenic, Tettegouche, and George H. Crosby-Manitou

State Parks. More than 146 species were documented in the red pine forest and 62 in the northern hardwood-conifer forest (Leacock 1997). Collections in St. Louis County, where this study was conducted, have been especially limited. There are only 31 specimens of macrofungi (18 species in six genera) recorded for St. Louis County in the University of

Minnesota Herbarium's Online Fungal Database (2002). The database contains only approximately 60% of the total fungi housed in the herbarium; however, it is quite probable that the database reflects the true state of knowledge for St. Louis County (D. McLaughlin, personal communication).

There is no fungal Red Data list for Minnesota, although there are six macrofungal species listed on the state Endangered, Threatened, and Special Concern Species List

(Minnesota Natural Heritage and Nongame Research Program 2002). There are no fungi included on the Region 9 Regional Forester Sensitive Species List which is used by the

Superior National Forest in northeastern Minnesota (USDA Forest Service 2000).

Although habitat conservation is typically the best way to conserve fungal diversity

(Staley 1997; Courtecuisse 2001), it is not always enough (Hawksworth 1992). Management practices and activities that maintain the habitat but potentially decrease diversity through soil disturbance must also be considered (Hawksworth 1992; Allison 2001). Prescribed burning, one such management practice, is used by land managers in northeastern Minnesota to mimic natural fire disturbance regimes, reduce ladder fuels and fuel loading, and create a suitable seed bed for red pine (Pinus resinosa Ait.) and eastern white pine (P. strobus L.). 9

Prescribed burning can increase the diversity of some fungi, particularly the carbinicolous fungi (Petersen 1970; Veerkamp 1998), or it can decrease the diversity of others, especially the mycorrhizal fungi (Stendell et al. 1999). Changes in species diversity can be short-lived (e.g., 1-4 years), as in the case of the relatively ephemeral carbiniocolous fungi (Petersen 1970). Most studies investigating the effects of prescribed burning on ectomycorrhizal fungi primarily focus on the immediate to short-term effects (Palmer et al.

1994; Stendell et al. 1999). Long-term effects of fire on macrofungal diversity in jack pine stands were investigated by Visser (1995), but the fire was a stand-replacing wildfire. The underlying, unavoidable conflict is that any type of management regime or disturbance will favor some species and inhibit others (Hobbs and Huenneke 1992).

Legal challenges to the development and implementation of forest management plans greatly shape the directions for managing public forests and have changed the focus from an emphasis on timber extraction to ecosystem management (Molina et al. 2001 ). If land managers are going to take an ecosystem approach to forest management, they must consider fungi. Fungi play significant ecosystem roles through nutrient cycling, food webs, forest diseases, and mutualisms, which influence seedling survival, tree growth, and overall forest health (Molina et al. 2001 ). To understand the biological and functional diversity of forest fungi, and before land managers in Minnesota can incorporate fungi into their management and conservation plans, fundamental information on the fungal species within the forest community and an understanding of the impacts of natural and anthropogenic disturbance must first be obtained.

One goal of this study was to obtain baseline quantitative data on species abundance, species richness, and fruiting phenology of the epigeous macro fungal community in northeastern Minnesota. A comprehensive picture of the fungal communities is needed as a basis for detecting changes in those communities. This study is an initial step towards such a foundation. Another goal was to investigate potential species composition differences in the fungal community following prescribed fire.

Study site and methods

Study site

The study was conducted in northeastern Minnesota in the Kawishiwi Ranger District of the Superior National Forest. The study site was 22.5 km north of Ely, near the Hegman lakes (48°02"N, 91 °55''W) at an elevation of 511 m (Fig. 1). The region, part of the Canadian

Shield (Ojakangas 1982), is characterized by Pre-Cambrian bedrock with a thin cover of glacial drift (USDA Forest Service 1998). The interagency National Hierarchical Ecological

Classification System refers to the local area surrounding the study site as the Trout Lake-

Indian Sioux Ground Moraine Landtype Association (212 La15) (USDA Forest Service

1998). This landtype association is characterized by a rolling to hilly ground moraine on top of Vermilion granite, and bedrock outcroppings are abundant. Upland soils are primarily shallow, well-drained Inceptisols derived from loamy till. Nutrient status is medium to low depending on depth of material to bedrock. Slopes at the study site range from 6 to 18 %.

The climate of northeastern Minnesota is continental, with short, warm, humid summers and long, cold, dry winters (Heinselman 1996). The mean annual precipitation is

711 mm, 64% of which occurs from May through September (Baker et al. 1967). The frost- free season lasts approximately 96 days, with the last spring frost occurring in early June and 11

I I I I I I I /\ I I \ I .,I \ I ,, \ I .,, \ I ' I I I I I Minnesota I I I I I I I I I I 3 I I :I /: /: I / I I / I \ \ ,' / \ I \, , __ .,. .,,' I ,' I I I I B control unit 4 plot ,/ I I ,' 2 I control unit 5 plot I I m I I I I gravel pit ',,, X .. '1 ...... I I ------burned unit boundary , I

I ---- BWCAW boundary «rlA""~L •••• •·············footpath Echo Trail I - - (County Road 116) Echo Trail

Fig. 1. Study site. (A) Location of site in Minnesota. (B) Study site showing the location of the burned units (Units I, 2, and 3) and the plots of the control units (Units 4 and 5) in relation to the Boundary Waters Canoe Area Wilderness (BWCAW). 12 the first autumn frost occurring in early September (Heinselman 1996). The lowest temperatures during the growing season occur in May, which has an average low of 4 °C and an average high of 18 °C. The highest temperatures occur in July, which has an average low of 13 °C and an average high of 26 °C.

The study site was a combination of two forest types: a near-boreal spruce-fir-aspen- birch forest and a Great Lakes red and white pine forest (Frelich 1998). The dominant tree species in the study plots were balsam fir (Abies balsamea (L) Mill.), red maple (Acer rubrum L.), paper birch (Betula papyrifera Marsh.), black spruce (Picea mariana (Mill.)

B.S.P.), red pine, eastern white pine, bigtooth aspen (Populus grandidentata Michx.), and quaking aspen (P. tremuloides Michx.). The ground vegetation was dominated by large- leaved aster (Aster macrophyllus L.), blue-bead lily ( Clintonia borealis (Ait.) Raf.), bunchberry (Cornus canadensis L.), wintergreen (Gaultheria procumbens L.), clubmosses

(Lycopodium spp.), Canada mayflower (Maianthemum canadense Desf.), bracken fern

(Pteridium aquilinum (L.) Kuhn), starflower (Trientalis borealis Raf.), blueberry (Vaccinium angustifolium Ait.), and various moss species. Each plot also included other plants that were less ubiquitous. Nomenclature of vascular plants follows Ownbey and Morley (1991).

The site was chosen because the Superior National Forest had conducted prescribed burns on three previously delineated units (Fig. 1). Unit 1 (30 ha) was burned in 1996, Unit

2 (70 ha) in 1997, and Unit 3 (32 ha) in 1999. All three burns were conducted in late May.

Three permanent plots, each measuring 10m x 10m square, were randomly located and established within each of the burned units in the summer of 1999, for a total of nine burn treatment plots. Also in 1999, six control plots of the same size were established in the surrounding unburned forest. The three unburned plots at the northern end of the study site 13 were collectively called Unit 4, while the three at the southern end were called Unit 5 (Fig.

1). Aside from the prescribed burning, the units experienced no other disturbance.

Sporocarp surveys

Fungal surveys were conducted twice in the early autumn of 1999. Fungi were surveyed on 12 and 11 occasions in 2000 and 2001, respectively, for a total of 25 surveys during the three-year study. In 2000 and 2001, surveys were conducted approximately every two weeks from mid-May until early October. Before and after these times the weather was not conducive to sporocarp production. Only true mushrooms and boletes growing on the soil surface and not on an obvious wood substrate were collected. During each survey, the date, species present, and number of sporocarps per species were noted for each plot to provide data on fruiting phenology, species richness, and species abundance (frequency and density). All sporocarps were picked and either saved for identification or documentation in a herbarium or discarded outside the plot to avoid double-counting during subsequent surveys. Harvesting sporocarps does not appear to have any negative short-term effects on subsequent fruiting (Norvell 1995; Pilz and Molina 2002).

Identification and ectomycorrhizal determination

Most sporocarps were identified in the lab following collecting trips. Specimens were dried on screens indoors without a heat source. Voucher specimens and supporting information are housed in Iowa State University's Ada Hayden Herbarium (ISC).

Nomenclature of fungi follows several sources (Table 1). Due to the high density of trees, it was extremely difficult to determine mycorrhizal associations between trees and sporocarps in the field. The fungus species were later categorized as ectomycorrhizal or non- mycorrhizal based on published literature and regardless of the geographical location of the 14

study (Table 1). Ectomycorrhizal status was classified as unknown if the genus of the tax on

was not positively determined, if there were conflicting reports in the literature about the

ecological role of the species, or if no information was found in the literature for the

particular species/genus (Table 1).

Data analysis

Three-year total data from the three plots within each unit were pooled, resulting in a

sample size of three burned units and two unburned units. Unit means were analyzed using a

t-test statistic (a= 0.05) to evaluate differences in species richness, ectomycorrhizal species richness, percent ectomycorrhizal species, and sporocarp abundance between the burned and unburned units. Differences between ectomycorrhizal and non-ectomycorrhizal species richness for all units combined were evaluated using a paired t-test (a= 0.5).

Results and discussion

Species diversity

After three seasons of sampling, 179 epigeous macrofungi, representing 33 genera and four orders (Agaricales, Boletales, Cortinariales, Russulales), were documented at the study site (Table 1). All of the 179 taxa were sufficiently distinctive to be classified as unique species despite the fact that positive species identifications were not always possible.

Identifications were made to the species or genus level for 156 of the 179 taxa collected, although not all were definitive. Twenty-three collections could not be identified at all and were categorized as "unknown". They are differentiated by their collection number (Table 15

1). The majority of the unknown species were collected in 1999 (Table 1) when the collection and preservation procedures were still being developed.

Of the 179 taxa, 167 (93 % ) were true mushrooms and 12 (7 % ) were boletes. Because boletes are ectomycorrhizal associates with many of the tree species present at the study site

(Molina et al. 1992), the small number of boletes collected was surprising. Conspicuously absent was Boletus edulis Bull., which was not documented inside the plots or even seen in the vicinity, yet it is frequently collected in the area by mushroom hunters.

Ectomycorrhizal or non-mycorrhizal status was determined for 149 of the 179 taxa

(Table 1). Of the 149 determined species, 68% (102 species) were ectomycorrhizal and 32%

(47 species) were non-mycorrhizal (Table 1). Exclusion of the undetermined taxa from these calculations could bias the results slightly if the 30 taxa had a different ratio of ectomycorrhizal to non-mycorrhizal species, but there is no reason to assume any difference.

Because all the dominant tree species in the boreal and temperate biomes form ectomycorrhizal associations that produce epigeous sporocarps (Molina et al. 1992; Vogt et al. 1992), it was expected that the majority of the fungal species collected at the study site would be ectomycorrhizal. When ectomycorrhizal species richness was compared to non- mycorrhizal species richness at the study site, the results showed more ectomycorrhizal species (unit x = 37) than non-mycorrhizal species (unit x = 17) within all the study units

(P=0.004). Bills et al. (1986) in West Virginia and Villeneuve et al. (1989) in Quebec also found coniferous forests to be dominated by ectomycorrhizal rather than saprobic fungi.

Approximately half of all the identified taxa were from just four genera. Russula (25 species), Cortinarius (23 species), Lactarius (11 species), and Amanita (9 species) were the most diverse groups in terms of species richness (Table 1). Other authors have also reported Table 1. Ectomycorrhizal status of epigeous fleshy fungi present on burned and/or control units in the Superior National Forest, Minnesota, 1999-2001. Species Bum Control ECM ECM Status Source Units Units Status a Boletales 2 Boletus subtomentosus L. b X X ECM Agerer 1987-1998 Boletus sp. (#313) c X ECM** Singer 1986 Chroogomphus corallinus O.K. Miller & Watl.5 X ECM** Singer 1986 Leccinum aurantiacum (Bull. ex St. Am.) S. F. Gray 10 X ECM* Bjurman and Fries 1984 Suillus granulatus (L.:Fr.) Roussel5 X X ECM Miller et al. 1983 Su illus pictus (Peck) A.H. Sm. & Theirs 10 X X ECM Palm and Stewart 1984

Suillus placidus (Bonord.) Sing.5 X ECM Miller et al. 1983 Suillus subalutaceous (A.H. Sm. & Thiers) A.H. Sm. & Thiers 1 X X ECM** Dahlberg and Finlay 1999 Suillus sp. (#K) X ECM** Singer 1986 Suillus sp. (#Z) X ECM** Singer 1986 Suillus sp. (#37) X ECM** Singer 1986 Suillus sp. (?) (#Y) X X ECM** Singer 1986 Ty lopilus felleus (Bull.:Fr.) Karst.5 X X ECM Jonsson et al. 1999 a Key to the specific ectomycorrhizal status of each species: ECM Ectomycorrhizal association experimentally determined. ECM* Putative~ association not experimentally determined, but species mentioned in the literature as ectomycorrhizal. ECM** Putative ~ species not mentioned in the literature, but genus predominately or exclusively ectomycorrhizal. NM Species mentioned in the literature as having a non-mycorrhizal ecological role. NMt Species in the genus are non-mycorrhizal according to Molina et al. ( 1992) and Singer ( 1986). ? Unknown status~ either conflicting reports in the literature, or genus known to have both mycorrhizal and non-mycorrhizal species, or species not identified sufficiently to determine an ecological role. b Nomenclature sources: 1. Bessette et al. 1997 5. Hansen and Knudsen 1992 9. Kibby and Fatto 1990 2. Bessette et al. 2000 6. Hesler and Smith 1963 10. Moser 1983 3. Brietenbach and Kranzlin 1991 7. Homola and Czapowskyj 1981 11 . Phillips 1981 4. Breitenbach and Kranzlin 2000 8. Kauffinan 1918 12. Phillips 1991 1--' 0\ c Collection number Table 1. (Continued) Species Bum Control ECM ECM Status Source Units Units Status Agaricales Amanita ceciliae (Berk. & Br.) Bas5 X ECM* Miller 1982 Amanita flavoconia Atk. 1 X ECM Godbout and Fortin 1985 Amanita fulva (Schaeff.) Pers. 5 X ECM Dahlberg et al. 1997 Amanitafulva (?) (Schaeff.) Pers.5 X ECM** Singer 1986 Amanita porphyria (Alb. & Schw.:Fr.) Mlady5 X ECM Dahlberg et al. 1997

Amanita vaginata (Bull. ex Fr.) Quel. 10 X ECM Peter et al. 2001 Amanita virosa (Fr.) G. Bertol.5 X X ECM* Salo 1993 Amanita sp. (#R) X ECM** Singer 1986 Amanita sp. (#206) X ECM** Singer 1986 Armillaria mellea complex (Vahl:Fr.) Kumm. 5 X X NM Salemi et al. 2001

Chrysomphalina sp. (#14) X NMt Singer 1986 Clitocybe clavipes (Pers.:Fr.) Kumm. 5 X X NM Chakravarty et al. 1999 Clitocybe sinopicoides Peck 11 X NMt Molina et al. 1992 Clitocybe vilescens (?) Peck8 X NMt Molina et al. 1992 Clitocybe sp. (#213) X NMt Singer 1986

Collybia butyracea (Bull.:Fr.) Kumm.5 X X NMt Singer 1986 Collybia confluens (Pers.:Fr.) Kumm. 5 X X NM Salo 1993 Collybia dryophila (Bull.:Fr.) Kumm. 5 X X NM Salemi et al. 2001 Collybia luteifolia Gill.3 X X NMt Molina et al. 1992 Collybia maculata (Alb. & Schw.:Fr.) Kumm. 5 X NM Salo 1993

Collybia proxilla (Fr.) Gill. 5 X NMt Molina et al. 1992 Coprinellus angulatus Peck 10 X NMt Molina et al. 1992 Entoloma majaloides Orton 10 X ? Hygrocybe acutoconica var. microspora (?) Hesler & A.H. Sm. 6 X NMt Singer 1986 5 Hygrocybe conica (Scop.:Fr.) Kumm. var. conica X NMt Singer 1986 >--' -..) Table 1. (Continued) Species Burn Control ECM ECM Status Source Units Units Status Hygrocybe laeta (Pers.:Fr.) Kumm. 5 X X NMt Singer 1986 Hygrocybe marginata Peck var. marginata6 X X NMt Singer 1986 Hygrocybe miniata (Fr.) Kumm.5 X X NMt Singer 1986 Hygrocybe miniata (?) (Fr.) Kumm. 5 X NMt Singer 1986 Hygrocybe miniata (Berk. & Br.) E. Arnolds var. mollis3 (?) X X NMt Singer 1986

Hygrophorus angustifolius (Murr.) Hesler & A.H. Sm.6 X X ECM** Homola and Czapowskyj 1985 Hygrophorus camarophyllus (Alb. & Schw.:Fr.) Dumee, Grandjean & Maire5 X X ECM* Salo 1993 Hygrophorus russula (Scop.:Fr.) Quel.5 X ECM* Salemi et al. 2001 Hygrophorus sp. (?) (#195) X ? Hygrophorus sp. (?) (#338) X ?

Hypholomafasciculare (Huds. :Fr.) Kumm.5 X NM Verhagen et al. 1998 Hypholomafasciculare (?) (Huds.:Fr.) Kumm. 5 X NMt Singer 1986 Laccaria bicolor (Maire) Orton5 X ECM Sung-Jae et al. 1998 Laccaria laccata (Scop.:Fr.) Berk. & Br. 5 X X ECM Godbold et al. 1998 Leptoniaformosus (Fr.) Quel. 10 X NMt Singer 1986

leptonia fulvus (?) (Orton) Mos. '0 X NMt Singer 1986 leptonia sericellum (Bull. ex Fr.) Kumm. 10 X NM Visser 1995 leptonia sp. (#43) X NMt Singer 1986 Leptonia sp. (#145) X NMt Singer 1986 Leptonia sp. (#199) X NMt Singer 1986

Marasmius androsaceus (L.:Fr.) Fr. 5 X NM Salemi et al. 2001 Marasmius sp. (#111) X NMt Singer 1986 Mycena epipterygia var. pelliculosa (?) (Fr.) Maas G. 5 X NM Salemi et al. 2001 Mycena griseoviridis A.H. Sm. 1 X NMt Singer 1986 Mycena griseoviridis (?) A.H. Sm. 1 X NMt Singer 1986 ...... 00 Table 1. (Continued) Species Bum Control ECM ECM Status Source Units Units Status Mycena sanguinolenta (Alb. & Schw.:Fr.) Kumm.5 X X NM Salemi et al. 2001 Mycena sp. (#102) X X NMt Singer 1986 Mycena sp. (#179) X NMt Singer 1986 Mycena sp. (#307) X NMt Singer 1986 Mycena sp. (#319) X NMt Singer 1986

Myxomphalia maura (Fr.) Hora5 X NM Norvell et al. 1994 Myxomphalia maura (?) (Fr.) Hora5 X NMt Singer 1986 Nolanea nitens (?) (Vel.) K. & R. 10 X NMt Singer 1986 Nolanea sp. (#116) X NMt Singer 1986 Pho/iota highlandensis (Peck) A.H. Sm. & Hesler5 X NMt Molina et al. 1992

Tricholoma apium (?) Schaeff.5 X X ECM** Singer 1986 Tricholoma columbetta (Fr.) Kumm. 5 X X ECM* Trappe 1962 Tricholomaflavovirens (Pers.: Fr.) Lundell5 X ECM Danielson 1984 Tricholoma imbricatum (?) (Fr.:Fr.) Kumm.5 X ECM** Singer 1986 Tricholoma magnivelare (Peck) Redhead 1 X ECM Gill et al. 2000

Tricholoma saponaceum (Fr.:Fr.) Kumm.5 X ECM Norkrans 1949 Tricholoma sejunctum (Sow.:Fr.) Quel.5 X X ECM Norkrans 1949 Tricholomopsis rutilans (Schaeff.:Fr.) Sing.5 X NM Salo 1993 Xeromphalina cauticinalis Ktihn. & Maire5 X X NMt Singer 1986

Cortinariales Cortinarius alboviolaceus (Pers.:Fr.) Fr. 5 X X ECM Godbout and Fortin 1985 Cortinarius armillatus (Fr.:Fr.) Fr.5 X ECM Agerer 1987-1998 Cortinarius bolaris (Pers.:Fr.) Fr.5 X X ECM Agerer 1987-1998 Cortinarius cinnamomeus (L.:Fr.) Fr.5 X ECM Agerer 1987-1998 Cortinarius leucopus (?) (Bull. ex Fr.) Fr. 10 X X ECM** Singer 1986 ...... \0 Table 1. (Continued) Species Bum Control ECM ECM Status Source Units Units Status Cortinarius lividoochraceus (?) (Berk.) Berk.5 X ECM* Salemi et al. 2001 Cortinarius multiformis var. coniferarum Mos. 10 X ECM* Dahlberg et al. 1997 Cortinarius obliquus Peck1 X X ECM Agerer 1987-1998 Cortinarius obtusus (?) (Fr.) Fr.5 X X ECM** Singer 1986 Cortinarius rubricosus (?) Fr. 10 X ECM** Singer 1986

Cortinarius semisanguineus (Fr.) Gill.5 X X ECM Agerer 1987-1998 Cortinarius sphaerosporus Peck1 X ECM** Singer 1986 Cortinarius subtorvus (?) Lamoure5 X X ECM** Singer 1986 Cortinarius tenebricus (?) Favre 10 X X ECM** Singer 1986 Cortinarius trivia/is Lange5 X ECM* Trappe 1962

Cortinarius sp. (#1/27) X X ECM** Singer 1986 Cortinarius sp. (# 15) X ECM** Singer 1986 Cortinarius sp. (#23b) X ECM** Singer 1986 Cortinarius sp. (# 152) X ECM** Singer 1986 Cortinarius sp. (#193) X ECM** Singer 1986

Cortinarius sp. (# 196) X ECM** Singer 1986 Cortinarius sp. (#317) X ECM** Singer 1986 Cortinarius sp. (#332) X ECM** Singer 1986 Cortinarius sp. (?) (#214) X ? Cortinarius sp. (?) (#318) X ?

Hebeloma crustuliniforme (Bull.) Quel.5 X X ECM Read 1998 Hebeloma sinapizans (?) (Paul.) Gill.5 X ECM Zambonelli et al. 2000 Hebeloma sp. (?) (#167) X ? lnocybe lacera (Fr.) Kumm.5 X ECM Cripps 1997 lnocybe leptophylla Atk.4 X ECM** Singer 1986 N 0 Table 1. (Continued) Species Bum Control ECM ECM Status Source Units Units Status lnocybe proximella (?) Karst. 4 X ECM** Singer 1986 Jnocybe sp. (#24/335) X ECM** Singer 1986 Jnocybe sp. (#25) X ECM** Singer 1986 Jnocybe sp. (#214) X ECM** Singer 1986 Naucoria sp. (#312) X ?

Phaeocollybia christinae (Fr.) Heims X NM Redhead and Malloch 1986 Phaeocollybiajennyae (Karst.) Heims X NM Redhead and Malloch 1986 Rozites caperatus (Pers.:Fr.) Karst.s X ECM Agerer 1999

Russulales Lactarius affinis var. affinis Peck1 X ECM Danielson 1984 Lactarius camphoratus (Bull.:Fr.) Fr. X X ECM Peter et al. 2001 Lactarius deceptivus Peck1 X X ECM* Bills et al. 1986 Lactarius hibbardae var. hibbardae Peck1 X X ECM Godbold et al. 1998 Lactarius lignyotus Fr.s X ECM Kraigher et al. 1995

Lactarius sordidus Peck 1 X ECM* Bills et al. 1986 Lactarius subvellereus var. subvellereus Peck7 X ECM* Homola and Czapowskyj 1981 Lactarius torminosus (Schaeff.:Fr.) Pers.s X ECM* Salo 1993 Lactarius uvidus (Fr.) Fr. var. uvidus 7 X ECM* Salemi et al. 2001 Lactarius vinaceorufescens A.H. Sm. 1 X X ECM* Bills et al. 1986

Lactarius sp. (#AA/E/26) X X ECM** Singer 1986 Russula bicolor (?) Burl.9 X ECM** Singer 1986 Russula brevipies Peck9 X X ECM Martinez-Amores et al. 1990/ l 99 l Russula brunneola Burl.9 X ECM** Singer 1986 Russula cyanoxantha (Schaeff.) Fr. f. cyanoxanthas X X ECM Beenken 2001

N Table 1. (Continued) Species Bum Control ECM ECM Status Source Units Units Status Russula decolorans (?) (Fr.) Fr.5 X X ECM** Singer 1986 Russula elaeodes (Bres.) Bon5 X ECM Agerer 1987-1998 Russulafaginea (?) Romagn.5 X ECM** Singer 1986 Russulafragilis (Pers.:Fr.) Fr.5 X X ECM Palmer et al. 1994 Russulafragrantissima Romagn.5 X X ECM* Doudrick et al. 1990

Russula incarnaticeps (?) Murrill9 X ECM** Singer 1986 Russula paludosa Britz.5 X ECM Dahlberg et al. 1997 Russula paludosa (?) Britz.5 (# 157) X ECM** Singer 1986 Russula paludosa (?) Britz.5 (#337) X X ECM** Singer 1986 Russula sanguinea (Bull.) Fr.5 X ECM Dufiabeitia et al. 1996

Russula sanguinea (?) (Bull.) Fr. 5 X X ECM** Singer 1986 Russula sp. (#J) X ECM** Singer 1986 Russula sp. (#38) X ECM** Singer 1986 Russula sp. (#171) X ECM** Singer 1986 Russula sp. (#200) X ECM** Singer 1986

Russula sp. (#314) X X ECM** Singer 1986 Russula sp. (#325) X ECM** Singer 1986 Russula sp. (#339) X ECM** Singer 1986 Russula sp. (#340) X ECM** Singer 1986 Russula sp. (#341) X ECM** Singer 1986 Russula sp. (#342) X ECM** Singer 1986

Unknown species: collection number and collection year # A/B, 1999 X X ? # C (Cortinarius sp. ?), 1999 X ? # P, 1999 X ? X N # Q, 1999 ? N # S, 1999 X ? Table 1. (Continued) Species Burn Control ECM ECM Status Source Units Units Status #U, 1999 X ? # V, 1999 X X ? # W, 1999 X ? # X, 1999 X ? # 16, 1999 X ? # 17, 1999 X ? # 18, 1999 X ? # 28, 1999 X ? # 32, 1999 X ? # 35 , 1999 X ? # 39, 1999 X ? # 42, 1999 X ? # 110, 2000 X ? # 113 , 2000 X ? # 153 , 2000 X ? # 201 , 2000 X ? # 215, 2000 X ? # 321 , 2001 X ?

N VJ 24 these to be amongst the most diverse genera in conifer-dominated ecosystems (Villeneuve et al. 1989; Salo 1993; Palmer et al. 1994; Dahlberg et al. 1997; Leacock 1997; Peter et al.

2001). Most genera were far less diverse, with 20 of the 33 genera represented by only one or two species (Table 1, Appendix B Table 1).

The burned units in the study area exhibited less species richness after three years of observation (unit x =51) than the unburned units (unit x =75) (pooled std. dev.=3.77,

P=0.006). There were also fewer ectomycorrhizal species present in the burned units (unit x = 32) than in the unburned units (unit x = 45) (pooled std. dev.=3.88, P=0.036). Prescribed burning always results in some fungal mortality because fire kills the fungal mycelium and ectomycorrhizae either directly or indirectly. The fungal mycelium and ectomycorrhizae in the organic horizons of the soil are killed directly when those horizons are consumed by the fire or when subsurface temperatures reach a lethal level. Ectomycorrhizae are killed indirectly when their host trees are killed by the fire. It is clear that the prescribed burning activities at the study site did impact the fungal community because the data show a decrease in species richness in the burned units.

Fire severity and intensity play a critical role in determining the degree to which fungi are affected. The less severe the burn, the fewer ectomycorrhizae are killed by organic layer consumption or lethal subsurface temperatures. Also, low intensity burns usually do not completely consume the duff and litter on the forest floor, leaving some substrate for the saprobic species. Although the intensities of the fires at the study site were not measured, the management objectives were successfully met, and the mortality of the large (DBH >40 cm) red and white pines was less than 20% (J. Hinds, personal communication). The data collected during this study do suggest that the bums, in general, did maintain a relatively low 25 intensity because, despite a reduction in ectomycorrhizal species richness in burned units, the percentage of ectomycorrhizal species present in the burned units (unit x = 68) was not less than the percentage in the unburned units (unit x = 70) (pooled std. dev.=0.067, P=0.779) If the burns had been consistently severe across the landscape, the majority of ectomycorrhizae likely would have been killed and not able to produce sporocarps; yet, this was not the case.

Despite the generally low intensity nature of the burns, the intensity did vary from unit to unit and also within each unit. However, it was assumed that the variation was random and that, by randomly locating the plots, the variation would not bias the results. In addition to the low intensity nature of the burns, many ectomycorrhizae might have also survived the fire due to their vertical distribution in the soil. Taylor and Bruns (1999) found that 60% of the ectomycorrhizal biomass on mature Pinus muricata roots occurred in the mineral soil, up to a sampling depth of 40 cm. It is possible that many of the ectomycorrhizal fungi in the burned units survived because they were distributed deeply in the soil.

There is a shortage of published research that monitors the long-term effects of prescribed burning on macrofungal species diversity in conifer-dominated ecosystems. Most studies investigated either the immediate to short-term effects of prescribed fire (Palmer et al.

1994; Stendell et al. 1999) or the short- to long-term effects of wildfire (Danielson 1984;

Visser 1995; Jonsson et al. 1999) on species diversity. In this study, the fungal communities were observed during a range of years, from the year of the burn (e.g., Unit 3 in 1999) to five years post-bum (e.g., Unit 1 in 2001). The pooled data suggest that five years following the low intensity prescribed burn, the fungal species richness was still not at pre-burn richness levels. There was an average of 51 species in a burned unit, while there was an average of 75 species in an unburned unit. Even, if burned richness levels were similar to pre-burn levels, 26 the species composition in burned and unburned areas would not necessarily be the same. Of the 179 taxa collected, 121 occurred in unburned areas, and 109 occurred in burned areas

(Table 1). Seventy species were unique to the unburned units, and 58 species were unique to the burned units (Table 1). There was a only a 29% overlap in species composition (51 species) between unburned and burned units (Table 1).

Some species occur specifically on burned-over areas and are called carbonicolous fungi (Dix and Webster 1995). Coprinellus angulatus, Leptonia sericellum, Marasmius androsaceus, Myxomphalia maura, Pholiota highlandensis, and Tricholomopsis rutilans have all been documented as occurring on burned ground (Smith and Hesler 1968; Peterson

1970). In this study, they were documented exclusively in burned units. There were no multi-taxa genera in this study that occurred exclusively in burned units (Table 1). However, all but two of the nine Amanita species occurred in burned units. All of the Amanita species found on P. muricata roots occurred primarily in the mineral soil at a depth greater than ~ 10 cm (Taylor and Bruns 1999). If the trend is not limited to Amanita species occurring with P. muricata, it might explain the ability of the Amanita species in the current study to survive and fruit in burned areas. Also, the Inocybes are considered "weedy" species that occur on disturbed sites (Kuyper 1986), and with the exception of one sporocarp, all of the Inocybes in this study were documented only in burned units (Table 1). There were also no multi-taxa genera that were limited to unburned sites. However, 16 of the 23 Cortinarius species occurred in Unit 4, which was not burned. Yet, Cortinarius is a dominant genus associated with aspen, in terms of species richness (Cripps 2001), and Unit 4 had the most aspen in the study area; therefore, the high occurrence of Cortinarius in Unit 4 might not have been due to the unburned nature of the unit. 27

Sporocarp abundance (density and frequency)

A total of 4,155 sporocarps was collected during the three-year study from the 1500 m2 total sample area, with 936, 2,240, and 979 sporocarps collected in 1999, 2000, and 2001, respectively (Appendix B Table 4). Total density of sporocarps at the study site was greatest in 2000, with 14,900 sporocarps/ha. This is similar to Leacock's (1997) estimate of 14,000-

16,000 sporocarps/ha for a Minnesota red pine forest. There were 6,500 sporocarps/ha in

2001. The 6,200 sporocarps/ha in 1999 is probably a low estimate as it represents data from only two sporocarp surveys for the year.

There was no difference between the sporocarp abundance on the burned units (unit x =223) and the unburned units (unit x =358) (pooled std. dev.=110, P=0.272). There was also no difference in the abundance of ectomycorrhizal sporocarps in the burned (unit x = 113) and unburned (unit x = 126) units (pooled std. dev.=33.7, P=0.719). There are many abiotic factors that influence sporocarp production, and while none is completely understood (Vogt 1992), it does appear that, in this case, prescribed fire did not negatively alter macrofungal productivity at the study site. O'Halloran et al. (1987) found that the numbers of macrofungal sporocarps did not differ between prescribed burned and unburned sites. Similarly, Palmer et al. (1994) found large numbers of ectomycorrhizal sporocarps on both burned and unburned sites immediately following a prescribed burn. Both groups suggested that prescribed fire did not appear to alter sporocarp production (O'Halloran et al.

1987; Palmer et al. 1994).

Of the 33 genera documented at the study site, the following five exhibited the most abundant sporocarp production and also accounted for more than half of all the sporocarps collected during the three-year study: Collybia (3,600 sporocarps/ha), Cortinarius (3,100 28

sporocarps/ha), Lactarius (2,200 sporocarps/ha), Laccaria (2,000 sporocarps/ha), and

Russula (1,900 sporocarps/ha) (Appendix B Table 2). Within those genera, one or two

species accounted for the majority of the sporocarps produced, with the exception of

Cortinarius. Six species of Collybia were documented, yet Collybia confluens and Collybia

dryophila together represented 91 % (3,300 sporocarps/ha) of the Collybia sporocarps.

Lactarius vinaceorufescens accounted for half of the Lactarius sporocarps (1,100

sporocarps/ha). All but two of the Laccaria sporocarps were Laccaria laccata (2,000

·sporocarps/ha). Russula fragi-lis accounted for over half of the sporocarps in that genus ··

(1,200 sporocarps/ha). With two species, Laccaria was not a diverse genus, yet it was a very

abundant genus in terms of sporocarp production. Leacock (1997), Dahlberg et al. (1997),

and Peter et al. (2001) found the same genera to be amongst the most productive in terms of

sporocarp abundance in coniferous forest ecosystems.

The following three species came from minimally diverse genera, but were very

abundant: Hypholoma fasciculare (1,800 sporocarps/ha), P. highlandensis (1,800

sporocarps/ha), and Armillaria mellea complex (1,100 sporocarps/ha). Hygrocybe (1,300

sporocarps/ha, 7 species) and Mycena (1,900 sporocarps/ha, 8 species) were both abundant

and relatively diverse genera (Appendix B Tables 1 and 2). It is interesting to note that many

of the highly productive, but less diverse, genera were non-mycorrhizal genera.

Species frequency varied widely across the landscape as only the following seven

species were documented on all five units: A. mellea complex, C. dryophila, Cortinarius

leucopus (?), L. laccata, Lactarius sp. (#AA/E/26), L. vinaceorufescens, and Mycena sp.

(#102). The majority of species (119, 66%) were documented on only one unit. Patchy and

uneven sporocarp distribution appears to be common in forest ecosystems. Peter et al. (2001) 29 found that nearly half of the species in a Picea abies forest in Switzerland were found in only one or two subplots. Also, more than half of the species in a P. abies forest in Sweden were documented in a single plot (Jonsson et al. 2000).

Fruiting phenology and annual variation

The fruiting season started in mid-May and lasted until early October. Sporocarp production peaked in September (Fig. 2). Leacock (1997) also found that sporocarp production in north central Minnesota was most abundant in September. Only 12 species fruited in May or June. The genera Hygrophorus, Phaeocollybia, Rozites, and Tricholoma fruited exclusively in September or October (Appendix B Table 3). Only one species,

Pholiota highlandensis, produced sporocarps every month of the growing season.

Of the 179 taxa collected during the three-year study, the numbers of species collected in 1999, 2000, and 2001 were 62 (35%), 113 (63%), and 85 (47%), respectively

(Appendix B Table 4 ). There was not much overlap, and fruiting of specific taxa was very inconsistent from year to year. Of the three-year taxa total, only 16 species (9%) fruited during every year of the study, while 114 species (64%) appeared in only one of the years

(Appendix B Table 4). Similarly, Dahlberg et al. (1997) found that nearly 50% of the macrofungal species in a Swedish, old-growth P. abies forest occurred in only one year of the six-year study. Ectomycorrhizal fungi appear to fruit less regularly from year to year than do saprobic fungi (Villeneuve et al. 1989), which makes it necessary to sample more than one year for an accurate community concept.

There were some differences in the seasonal fruiting patterns of species unique to the burned units and those unique to the unburned units (Fig. 3). Species in the burned units produced more sporocarps in the spring, early summer, and late fall than did species in the 140

120

100 fl) QJ "cj QJ C. fl) 80 (.- 0 i... QJ .Q 60 E z::: 40

0 May June July August September October Month

Fig. 2. Total number of epigeous macrofungal species that were fruiting each month of the growing season, within the study site, and over the course of the study (1999-2001). The study site, a mixture of near-boreal and Great Lakes red and white pine forests, is located in the Superior National Forest, Minnesota.

VJ 0 100

b() .5 70 ·a c!:- fl) 60 ·u 50 c..fl) '- 0 40 C - Burned Unburned C.J D I,., 30 I

0 May June July August September October

Month

Fig. 3. Percent of epigeous macrofungal species fruiting exclusively on burned and unburned units during each month of the growing season. Percentages reflect combined data from the three year study, 1999-2001. The study site, a mixture of near-boreal and Great Lakes red and white pine forests, is located in the Superior National Forest, Minnesota.

u.) J,,,-,l 32 unburned units (Fig. 3). The species in burned units exhibited a fruiting pattern that was relatively constant throughout the growing season, and the pattern lacked the extreme spike in production that the species in unburned units produced in September, although fruiting was still the most productive in September (Fig. 3).

More than half of the 12 species that fruited in May and June occurred only in burned units: P. highlandensis, C. angulatus, lnocybe lacera, Lactarius lignyotus, Marasmius androsaceus, Mycena sp. (#307), and M. maura (Appendix B Table 5). Peterson (1970) found that fireplace fungi were able to produce sporocarps throughout the spring, summer, and autumn providing there was sufficient soil moisture. The conditions of burned soil in the spring would seem to favor sporocarp production (Vogt et al. 1992). Burned soil has the potential to be warmer than unburned soil due to increased solar heating of the open, exposed soil (Neary et al. 1999), and soil moisture content is typically high in the spring as a result of the snowmelt (Heinselman 1996). If present, those conditions might explain why the burned units in the current study exhibited a higher degree of fruiting in the spring and early summer months than the unburned units. Fewer species fruited in August and September in the burned units than in the unburned units (Fig. 3). The open exposed soil, the same trait that was beneficial in the spring, is no longer beneficial in the summer when evapotranspiration rates increase, resulting in low soil moisture and unfavorable fruiting conditions (Peterson

1970; Neary et al. 1999). Peterson (1970) suggested that the ability to fruit during any period with favorable temperature and moisture conditions must be advantageous for fungi that occur exclusively or primarily in the relatively ephemeral habitat of a burned site.

To summarize, 179 taxa were found at the study site during three seasons of sampling. The dominant genera, in terms of species richness and sporocarp abundance, were 33 consistent with the observations of many studies conducted in conifer-dominated forests.

The species richness and ectomycorrhizal species richness were lower on burned units compared to unburned units. However, the percentage of ectomycorrhizal species was not different between burned and unburned units, which suggests either that the bums were of sufficiently low intensity, or that the ectomycorrhizal species were deep enough in the soil to avoid lethal subsurface temperatures, or that there was a combination of the two. These baseline data on the epigeous macrofungal community should serve as a foundation on which future work can be based, in order to understand even more completely the fungal community of the near-boreal/Great Lakes pine forests of northeastern Minnesota.

Acknowledgments

Very special thanks to Tom Yankowiak for helping conduct the fungal surveys.

Sincere gratitude to Anna Gardner for creating the study site map. Thanks to Jim Hinds and

Robert Kari, USDA Forest Service, and Dan Hanson, Minnesota Department of Natural

Resources, for their technical expertise. Many thanks to David McLaughlin, curator of the

University of Minnesota Fungal Herbarium, for his valuable input. I sincerely appreciate the editorial comments and support of Lois H. Tiffany, J. Michael Kelly, and Thomas Jurik at

Iowa State University. 34

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CHAPTER 3. SHORT-TERM EFFECTS OF PRESCRIBED BURNING ON THE ECTOMYCORRHIZAL FUNGAL COMMUNITY IN A NEAR-BOREAL/ GREAT LAKES PINE FOREST IN MINNESOTA.

A paper submitted to the Canadian Journal of Forest Research

Erin J. Heep

Abstract

Prescribed burning is a common management practice used by public lands managers, yet there are no published data on the effects of prescribed burning on the macrofungal community in northern Minnesota coniferous forests. I determined the short-term effects of prescribed burning on the epigeous, ectomycorrhizal, macrofungal community in a near- boreal/Great Lakes pine forest. Sporocarps were surveyed in 1999, 2000, and 2001 in the

Superior National Forest to determine ectomycorrhizal sporocarp richness and abundance.

Of the 149 taxa identified to genus or species, 102 (68%) were ectomycorrhizal and 47 (32%) were non-mycorrhizal. Ectomycorrhizal species richness was lower in burned units than in unburned units, as was overall species richness. The percent of ectomycorrhizal species did not differ between treatments. There was no difference in the overall or ectomycorrhizal sporocarp abundance between burned and unburned units. Low intensity prescribed burning consumes much of the organic portion of the soil and with it, the ectomycorrhizae. Yet, the ectomycorrhizae were not completely consumed by the fire in this study. It is suggested that they either survived in the unburned organic matter or were deep enough in the soil to be unaffected by temperatures that are lethal only in the top few centimeters of the soil. 43

Introduction

Approximately 13% (1.4 million ha) of northern Minnesota's forests (10.9 million ha total) was dominated by red pine (Pinus resinosa Ait.) and eastern white pine (P. strobus L.) prior to European settlement from 1850 to 1900 (Frelich 1998). Extensive logging followed by high-intensity slash fires decimated the Great Lakes pine forests (Rusterholz 1990;

Heinselman 1996; Haapoja 1997), and today they account for only 3% (0.2 million ha) of northern Minnesota's forest types (6.7 million ha total) (Frelich 1998). Development and preservation of red and white pine forests is favored by a fire regime of infrequent crown fires (150-250 year intervals) and frequent surface fires (10-50 year intervals) (Heinselman

1996). Due to a lack of frequent surface fires, red and white pines have not been consistently regenerating and are being replaced by more shade tolerant species such as balsam fir (Abies balsamea (L.) Mill.), red maple (Acer rubrum L.), paper birch (Betula papyrifera Marsh.), and black spruce (Picea mariana (Mill.) B.S.P.) (Wendt and Coffin 1988; Heinselman 1996;

Frelich 1998).

Prescribed burning as a tool for land management purposes has increased substantially in the past two decades (Neary et al. 1999). Although fire is a natural process and was instrumental in shaping the forests we have today (Ahlgren and Ahlgren 1960;

Ahlgren 1974), fire can also have deleterious short- to long-term effects on the ecosystem through changes in below-ground components and processes (Neary et al. 1999). It is primarily the fire severity that determines whether the effects of fire will be beneficial or harmful (Neary et al. 1999), although frequency, intensity, and depth of heat pulse also play a role (Staddon et al. 1996). Prescribed bums are usually not severe fires, so one might assume that the effects would not be harmful. However, even low intensity surface fires can, under 44 certain conditions, completely consume the organic layer (Frandsen and Ryan 1986; Hartford and Frandsen 1992), and temperatures can reach biologically lethal levels within the top few centimeters of mineral soil (Hartford and Frandsen 1992; Oliveira et al. 1997; Neary et al.

1999). The majority of ectomycorrhizae are concentrated in the organic horizons of the soil and top few centimeters of the mineral soil (e.g., Harvey et al. 1976; Buchholz and Motto

1981; Persson 1983). Thus, prescribed fire can kill ectomycorrhizae directly by consuming the organic layers of the soil and by raising sub-surface temperatures to a lethal level. For example, Dahlberg et al. (2001) found that four months after a slight-bum treatment, mycorrhizae occurred only in the mineral soil of a Swedish boreal forest. This suggests that mycorrhizae in the organic layers of the soil would be killed even during a low severity natural or prescribed burn (Dahlberg et al. 2001). Some ectomycorrhizae may be killed indirectly by fire when the flow of carbohydrates from the tree to the fungus ceases as a result of tree mortality after a prescribed burn. Hacskaylo (1973) suggested that without the host-supplied substrate, ectomycorrhizal fungi would not persist long. Harvey et al. ( 1980) demonstrated that root systems could not sustain mycorrhizae beyond ~ 2 months after tree mortality. Therefore, it is possible that even a low intensity prescribed burn could have harmful effects on the ectomycorrhizal fungal community.

Some studies have determined the effects of prescribed fire or wildfire on the ectomycorrhizal community by surveying the epigeous sporocarps (O'Halloran et al. 1987;

Palmer et al. 1994; Visser 1995; Jonsson et al. 1999; Dahlberg et al. 2001), whereas others have surveyed the below-ground ectomycorrhizal community by analyzing the ectomycorrhizae present on the fine-root tips of trees (Visser 1995; Jonsson et al. 1999;

Stendell et al. 1999; Dahlberg et al. 2001; Mah et al. 2001). Studies have been conducted in 45

Europe (Visser 1995; Torres and Honrubia 1997; Jonsson et al. 1999; Dahlberg et al. 2001), western North America (Horton et al. 1998; Amaranthus and Trappe 1993; Stendell et al.

1999; Mah et al. 2001), and the southern (O'Halloran et al. 1987; Palmer et al. 1994) and eastern (Buchholz and Gallagher 1982) United States, but not the north central United States.

The Superior National Forest in northeastern Minnesota has identified a section of

Great Lakes pine forest near the Hegman Lakes in the Boundary Waters Canoe Area

Wilderness (Fig. 1) that has largely succeeded to a near-boreal spruce-fir-aspen-birch forest.

In an attempt to return this area to its former forest cover type, prescribed burning was used to mimic natural surface fires. The prescribed burning plan was implemented with the assumption that the fire would consume the accumulated pine needles, litter, and upper humus layers and kill the shrubs and invading shade-tolerant species. Red and white pine seedlings could then become established, and the Great Lakes pine forest in that area would be restored. Studies that determine the effects of prescribed fire on the ectomycorrhizal fungal community in that forest type are needed because it is difficult to estimate the ecological impacts of fire on one ecosystem based on studies conducted in another (Herr et al. 1994 ), because prescribed burning will continue to be used as a management tool in the

Superior National Forest, and because ectomycorrhizal fungi play a critical role in forest ecosystems.

The goal of this study was to evaluate the short-term effects of prescribed burning on ectomycorrhizal fungi in a near-boreal/Great Lakes pine forest by comparing above-ground ectomycorrhizal species richness and abundance in burned and unburned stands. It was hypothesized that the response of ectomycorrhizal fungi to prescribed burning would be a 46 decrease in species richness and abundance resulting from organic layer consumption, lethal sub-surface temperatures, and some tree mortality.

Study site and methods

Study site

The study was conducted 22.5 km north of Ely, Minnesota, near the Hegman lakes

(48°02 "N, 91 °55 "W, elevation 511 m) in the Kawishiwi Ranger District of the Superior

National Forest (Fig. 1). The region, part of the Canadian Shield (Ojakangas 1982), is characterized by Pre-Cambrian bedrock with a thin cover of glacial drift (USDA Forest

Service 1998). The local area surrounding the study site is the Trout Lake-Indian Sioux

Ground Moraine Landtype Association (212 Lal5), according to the interagency National

Hierarchical Ecological Classification System (USDA Forest Service 1998). This landtype association is characterized by a rolling to hilly ground moraine on top of Vermilion granite.

Upland soils are primarily shallow, well-drained Inceptisols derived from loamy till, and bedrock outcroppings are abundant. Nutrient status is medium to low depending on depth of material to bedrock. Slopes at the study site range from 6 to 18%.

Climate is continental, with short, warm, humid summers and long, cold, dry winters

(Heinselman 1996). Mean annual precipitation is 711 mm, 64% of which occurs from May through September (Baker et al. 1967). There are approximately 96 frost-free days, with the last spring frost in early June and the first autumn frost in early September (Heinselman

1996). The coldest temperatures of the growing season are in May, which has an average low of 4 °C and an average high of 18 °C. The warmest growing season temperatures are in

July, which has an average low of 13 °C and an average high of 26 °C. 47

B t 0.5 km N

I I I I I I I I\ I I \ I I \ I ,,,,," \ ,' ,, I / Minnesota ,/ : / : : I I I I I 3 I I I / / I I I I / / I ,' ,' ,,,/ \, , __ ,,,. ,,, / 8 control unit 4 plot I

control unit 5 plot I/ 2 m I I gravel pit \ X ' , __ ------burned unit boundary ---- BWCAW boundary

•·············footpath m I Echo Trail I - - (County Road 116) Echo Trail

The study area was a mixture of two forest types: a near-boreal spruce-fir-aspen- birch forest and a Great Lakes red and white pine forest (Frelich, 1998). The dominant tree species in the study plots were balsam fir, red maple, paper birch, black spruce, red pine, eastern white pine, bigtooth aspen (Populus grandidentata Michx.), and quaking aspen (P. tremuloides Michx.). The ground vegetation was dominated by large-leaved aster (Aster macrophyllus L.), blue-bead lily ( Clintonia borealis (Ait.) Raf.), bunchberry ( Cornus canadensis L.), wintergreen (Gaultheria procumbens L.), clubmosses (Lycopodium spp.),

Canada mayflower (Maianthemum canadense Desf.), bracken fem (Pteridium aquilinum (L.)

Kuhn), starflower (Trientalis borealis Raf.), blueberry (Vaccinium angustifolium Ait.), and various moss species. Each plot also included other plants that were less ubiquitous.

Nomenclature of vascular plants follows Ownbey and Morley (1991).

The study site was chosen because the Superior National Forest had conducted prescribed bums on three previously delineated units (Fig. 1). Unit 1 (30 ha) was burned in

1996, Unit 2 (70 ha) in 1997, and Unit 3 (32 ha) in 1999. All units were burned in late May.

Three permanent plots, each measuring 1Om x 1Om square, were randomly located and established within the burned units in the summer of 1999, for a total of nine treatment plots.

Also in 1999, six control plots of the same size were established in the surrounding unburned forest. The three unburned plots at the northern end of the study site were collectively called

Unit 4, while the three at the southern end were called Unit 5 (Fig. 1). The units experienced no disturbance apart from the prescribed burning.

Sporocarp surveys and identification

Fungal surveys were conducted twice in the early autumn of 1999. In 2000 and 2001, surveys were conducted approximately every two weeks from mid-May until early October, 49 for a total of 12 and 11 surveys, respectively. True mushrooms and boletes were collected, but only those growing on the soil surface and not on any obvious wood substrate. During each survey, the date, species present, and number of sporocarps per species in each plot were recorded to provide data on fruiting phenology, species richness, and species abundance

(frequency and density). All sporocarps were picked to avoid double-counting during subsequent surveys and were either identified and discarded outside the plot, or saved for later identification and preservation in a herbarium. Harvesting sporocarps does not appear to have any negative short-term effects on subsequent fruiting (Norvell 1995; Pilz and

Molina 2002). Some identifications were made in the field, but the majority of sporocarps were identified later in the lab. Specimens were dried on screens indoors without a heat source. Voucher specimens, spore prints, photographs, and data forms are housed in Iowa

State University's Ada Hayden Herbarium (ISC). Nomenclature of fungi follows several sources (Chapter 2, Table 1).

Ectomycorrhizal determination

Due to the high density of trees, it was extremely difficult to determine mycorrhizal associations between trees and sporocarps in the field. After identification, the fungi were categorized as ectomycorrhizal or non-mycorrhizal based on existing literature, regardless of the geographical location of the published study. Ectomycorrhizal status was classified as unknown if the genus of the taxon was undetermined, if there were conflicting reports in the literature about the ecological role of the species, or if no information was found in the literature for the particular species/genus. 50

Data Analysis

Three-year total data from the three plots within each unit were pooled, resulting in a sample size of three burned units and two unburned units. Unit means were analyzed using a t-test statistic (a= 0.05) to evaluate differences in species richness, ectomycorrhizal species richness, percent ectomycorrhizal species, overall sporocarp abundance, and ectomycorrhizal sporocarp abundance between the burned and unburned units.

Results and discussion

After three seasons of sampling, 179 taxa were documented at the study site. Of the total taxa, 167 (63%) were true mushrooms and 12 (7%) were boletes. Identifications were made, to at least the genus level, for 156 taxa, although not all were definitive Twenty-three collections were not identified. Ectomycorrhizal status was determined for 149 of the 179 taxa. The 30 taxa that were not assigned an ectomycorrhizal status were the 28 unidentified species, five taxa for which the genus identification was uncertain, Naucoria sp. (#312), and

Entoloma majaloides Orton (Chapter 2 Table 1). Of the 149 taxa for which an ectomycorrhizal status was determined, 102 (68%) were ectomycorrhizal and 47 (32%) were non-mycorrhizal. Other researchers have also found that ectomycorrhizal fungi are the dominant ecological group in coniferous forests (Bills et al. 1996, Villeneuve et al. 1989).

The dominant genera at the study site, in terms of species richness, were Russula (25 species), Cortinarius (23 species), Lactarius (11 species), and Amanita (9 species). All four genera are considered ectomycorrhizal genera (Singer 1986). The dominant genera in terms of sporocarp abundance included ectomycorrhizal and non-mycorrhizal genera. The four previously mentioned genera , as well as Laccaria, were the abundant ectomycorrhizal 51 genera. Non-mycorrhizal genera that produced abundant sporocarps were Collybia,

Hypholoma, Mycena, and Pholiota. For a complete list of species documented at the study site, their mycorrhizal status, and their distribution amongst the burned and/or unburned units, see Table 1 in Chapter 2.

Three-year ectomycorrhizal species richness was lower in the burned units (unit

x = 32) than in the unburned units (unit x = 45) (pooled std. dev .=3.88, P=0.036). There were also fewer species in general in the burned units (unit x = 51) than in the unburned units

(unit x =75) (pooled std. dev.=3.77, P=0.006). Because ectomycorrhizal species are dominant at the study site, and because there were fewer species in general in the burned units, it is not surprising that there were also fewer ectomycorrhizal species in the burned units. Fewer ectomycorrhizal species in the burned units suggests that the fire killed, directly or indirectly, some ectomycorrhizae, or that the fire at least inflicted sufficient damage on the vegetative mycelia so that sporocarps were not produced.

Several studies have demonstrated that above-ground sporocarp surveys do not completely reflect the diversity and composition of the below-ground ectomycorrhizal fungal community because many important ectomycorrhizal species produce either inconspicuous epigeous sporocarps, hypogeous sporocarps, or no sporocarps, and not all sporocarps produced are true mushrooms or boletes (e.g., Danielson 1984; Gardes and Bruns 1996;

Dahlberg et al. 1997; Jonsson et al. 1999; Jonsson et al. 2000). When evidence that the vegetative body of the fungus exists underground as a mycorrhiza in healthy condition is obtained only with the presence of an epigeous basidiocarp, it is difficult to determine the full effects of fire on the entire ectomycorrhizal community. Nevertheless, sporocarp surveys are valuable and have advantages over below-ground surveys in many ways. For example, 52 extensive areas or large numbers of plots can be surveyed and with minimal interference to the fungi (Dahlberg et al. 1997; Peter et al. 2001). Also, most taxa can be identified to species, and sporocarp surveys allow the less-frequent and uncommon species to be detected

(Dahlberg et al. 1997; Dahlberg et al. 2001).

Realizing that the effects of fire on ectomycorrhizal fungi will not be completely understood through sporocarp surveys alone, the results of this study indicate that the fire killed sufficient ectomycorrhizae to significantly reduce the sporocarp richness in the burned units. Dahlberg et al. (2001) found that a hard-bum treatment killed all the mycorrhizae, whereas a slight-bum treatment reduced the mycorrhizae in a Swedish boreal forest, but left much of the mycorrhizal flora alive in the soil. The mycorrhizae remaining after the slight- bum treatment were encountered only in the mineral soil portion of the core samples, suggesting that mycorrhizae existing in the organic portions of the soil would normally be killed, regardless of fire severity (Dahlberg et al. 2001). Because the majority of fine-roots and mycorrhizae occur in the organic portions of the soil (Finer et al. 1997; Jonsson et al.

2000; Dahlberg et al. 2001 ), it is quite reasonable to assume that some or most mycorrhizae in the organic layers would be destroyed as fire consumes the forest floor. The results of this study support this conjecture, as there were fewer ectomycorrhizal species present in the burned units.

Even though there were fewer ectomycorrhizal present after the bums at the study site, there was no difference in the percentage of ectomycorrhizal species occurring in the burned units (unit x = 68) compared to the unburned units (unit x = 70) (pooled std. dev.=0.067, P=0.779). Those ectomycorrhizae that produce mushrooms and boletes were not so adversely affected by the fire as to lead to a significant decrease in the percentage of 53 ectomycorrhizal fungi in the burned units. In general, the intensity of the fires appears to have remained relatively low, as an insignificant percentage of ectomycorrhizae were killed, either directly or indirectly.

There was no difference between the burned and unburned units in terms of overall sporocarp abundance. The burned units (unit x = 223) did not differ from the unburned units

(unit x = 358) (pooled std. dev.=110, P=0.272) in the average number of sporocarps produced over three years in each unit at the study site. There was also no difference in the abundance of ectomycorrhizal sporocarps in the burned (unit x = 113) and unburned (unit

x = 126) units (pooled std. dev.=33.7, P=0.719). O'Halloran et al. (1987) also found no difference between prescribed burned and unburned sites in terms of sporocarp production in a Texas pine-hardwood forest. There are many factors, e.g., precipitation, that likely influence annual and spatial fruiting patterns of basidiocarps (Vogt et al. 1992). However, the results of this study indicate that fire did not negatively alter the abundance of sporocarps produced during the three-year study.

Of the 102 ectomycorrhizal species documented at the study site, 30% were found exclusively in the burned units, 35% exclusively in the unburned units, and 35% in both burned and unburned units. A similar pattern emerged in a Swedish boreal forest where

Dahlberg et al. (2001) found that 43% of species were unique to burned forest, 37% were unique to unburned forest, and 20% were found in both burned and unburned forest, indicating that fire can alter the near-term species composition of the ectomycorrhizal community.

Although there was no difference between the percentage of ectomycorrhizal species in burned and unburned units, there was a difference in species composition. Only 54 approximately one third (35%) of the species overlapped. It has been suggested that some ectomycorrhizal taxa prefer different soil depths (Taylor and Bruns 1999; Dahlberg et al.

2001; Peter et al. 2001). It is possible that, in this study, some of the ectomycorrhizal fungi found in the burned units were consistently associated with roots that were deeper in the soil.

It is possible that some of the ectomycorrhizal fungi found only in the unburned units had an affinity for the organic soil layers and were eliminated from the burned units as a result of the fire consuming the organic material. This assumes that the ectomycorrhizal fungi found only in unburned units would have probably occurred also in the burned units prior to the bum.

However, no pre-bum data exist for the burned units. In this study, six of the nine Amanita species collected occurred only in burned units, two occurred only in unburned units, and one species occurred in both types of units. Taylor and Bruns ( 1999) found that, of the Pinus muricata roots colonized by four Amanita species, the vast majority occurred in mineral soil at or deeper than ~ 10 cm. If the genus Amanita has a preference for greater depths, it might explain why so many Amanita species were found in burned units. Their deep distribution in the mineral soil would have given them an advantage over the shallower fungi during a bum.

This does not necessarily explain, however, why there were not equally high numbers of

Amanita species in the unburned units. Taylor and Bruns (1999) speculated that Amanita species were relatively weak vegetative competitors (compared to Russula species). If so, they would possibly have an advantage when species that formed mycorrhizae nearer the surface, or species that more aggressively colonized roots, were consumed by the fire.

Ectomycorrhizal fungi perform a valuable role in northern coniferous forest ecosystems by enhancing nutrient acquisition, drought tolerance, and pathogen resistance of their hosts (Peter et al. 2001). It is therefore important that forest managers who utilize 55 prescribed burning in their management practices understand, as much as possible, the potential effects of fire on the various components of the ecosystem, including the ectomycorrhizal fungal community. This study demonstrates that, while tentative conclusions can be made as to the effects of fire on ectomycorrhizal fungi, it is difficult to make concrete conclusions. While ectomycorrhizal species richness decreased immediately following a low intensity bum, many mycorrhizae survived and produced abundances and species richnesses that were no different from the unburned units. Despite the fact that species with a shallow distribution of mycelia would probably decrease after a bum

(Dahlberg at al. 2001), it appears that the ectomycorrhizal fungal community, in the long- term, would not generally be adversely affected by low intensity prescribed bums.

Acknowledgments

Very special thanks to Tom Yankowiak for helping conduct the fungal surveys.

Sincere thanks to Anna Gardner for creating the study site map. Many thanks to David

McLaughlin, curator of the University of Minnesota Fungal Herbarium, for his valuable input. Thanks to Jim Hinds and Robert Kari, USDA Forest Service, and Dan Hanson ,

Minnesota Department of Natural Resources, for their technical expertise. I sincerely appreciate the editorial comments and support of Lois H. Tiffany, J. Michael Kelly, and

Thomas Jurik at Iowa State University. 56

References

Ahlgren, I.F., and Ahlgren, C.E. 1960. Ecological effects of forest fires. Bot. Rev. 26: 483- 533.

Ahlgren, C.E. 1974. Effects of fires on temperate forests: north central United States. In Fire and ecosystems. Edited by T.T. Kozlowski and C.E. Ahlgren. Academic Press, New York, New York. pp. 195-223.

Amaranthus, M.P., and Trappe, J.M. 1993. Effects of erosion on ecto- and VA-mycorrhizal inoculum potential of soil following forest fire in southwest Oregon. Plant Soil, 150: 41- 49.

Baker, D.G., Haines, D.A., and Strub, J.H., Jr. 1967. Climate of Minnesota. V. Precipitation facts, normals, and extremes. Minn. Agric. Exp. Stn. Tech. Bull. 254. Minneapolis, Minnesota.

Buchholz, K., and Gallagher, M. 1982. Initial ectomycorrhizal density response to wildfire in the New Jersey pine barren plains. Bull. Torrey Bot. Club, 109: 396-400.

Buchholz, K., and Motto, H. 1981. Abundances and vertical distributions of mycorrhizae in plains and barrens forest soils from the New Jersey pine barrens. Bull. Torrey Bot. Club, 108: 268-271.

Dahlberg, A., Schimmel, J., Taylor, A.F.S., and Johannesson, H. 2001. Post-fire legacy of ectomycorrhizal fungal communities in the Swedish boreal forest in relation to fire severity and logging intensity. Biol. Conserv. 100: 151-161.

Danielson, R.M. 1984. Ectomycorrhizal associations in jack pine stands in northeastern Alberta. Can. J. Bot. 62: 932-939.

Finer, L., Messier, C., and de Grandpre, L. 1997. Fine-root dynamics in a mixed boreal conifer - broad-leafed forest stands at different successional stages after fire. Can. J. For. Res. 27: 304-314.

Frandsen, W.H., and Ryan, K.C. 1986. Soil moisture reduces belowground heat flux and soil temperatures under a burning fuel pile. Can. J. For. Res. 16: 244-248. 57

Frelich, L.E. 1998. Natural disturbance and variability of forested ecosystems in northern Minnesota. USDA Forest Service, Superior National Forest, Duluth, Minnesota.

Gardes, M., and Bruns, T.D. 1996. Communtiy structure of ectomycorrhizal fungi in a Pinus muricata forest: above- and below-ground views. Can. J. of Bot. 74: 1572-1583.

Haapoja, M.A. 1997. Renaissance for white pine. Minnesota Volunteer, 60: 10-19.

Hacskaylo, E. 1973. Carbohydrate physiology of ectomycorrhizae. In Ectomycorrhizae: their ecology and physiology. Edited by G.C. Marks and T.T. Kozlowski. Academic Press, New York, New York. pp. 207-230.

Hartford, R.A., and Frandsen, W.H. 1992. When it's hot, it's hot. .. or maybe it's not! (Surface flaming may not portend extensive soil heating). Int. J. Wildland Fire, 2: 139- 144.

Harvey, A.E., Larsen, M.J., and Jurgensen, M.F. 1976. Distribution of ectomycorrhizae in a mature douglas-fir/larch forest soil in western Montana. For. Sci. 22: 393-398.

Harvey, A.E. Jurgensen, M.F., and Larsen, M.J. 1980. Clearcut harvesting and ectomycorrhizae: survival of activity on residual roots and influence on a bordering forest stand in western Montana. Can. J. For. Res. 10: 300-303.

Heinselman, M.L. 1996. The boundary waters wilderness ecosystem. University of Minnesota Press, Minneapolis, Minnesota.

Herr, D.G., Duchesne, L.C., Tellier, R., McAlpine, R.S., and Peterson, R.L. 1994. Effect of prescribed burning on the ectomycorrhizal infectivity of a forest soil. Int. J. Wildland Fire, 4: 95-102.

Horton, T.R., Cazares, E., and Bruns, T.D. 1998. Ectomycorrhizal, vesicular-arbuscular and dark septate fungal colonization of bishop pine (Pinus muricata) seedlings in the first five months of growth after wildfire. Mycorrhiza, 8: 11-18.

Jonsson, L., Dahlberg, A., and Brandrud, T-E. 2000. Spatiotemporal distribution of an ectomycorrhizal community in an oligotrophic Swedish Picea abies forest subjected to experimental nitrogen: above- and below-ground views. For. Ecol. Manag. 132: 143-156.

Mah, K., Tackaberry, L.E., Egger, K.N., and Massicotte, H.B. 2001. The impacts of broadcast burning after clear-cutting on the diversity of ectomycorrhizal fungi associated with hybrid spruce seedlings in central British Columbia. Can. J. For. Res. 31: 224-235. 58

Neary, D.G., Klopatek, C.C., DeBano, L.F., and Ffolliott, P.F. 1999. Fire effects on belowground sustainability: a review and synthesis. For. Ecol. Manag. 122: 51-71.

Norvell, L.L. 1995. Loving the chanterelle to death? The ten-year Oregon chanterelle project. Mcllvainea, 12: 6-25.

O'Halloran, K.A., Blair, R.M., Alcaniz, R., and Hershel, F.M., Jr. 1987. Prescribed burning effects on production and nutrient composition of fleshy fungi. J. Wildl. Manag. 51: 258- 262.

Ojakangas, R.W., and Matsch, C.L. 1982. Minnesota's geology. University of Minnesota Press, Minneapolis, Minnesota.

Oliveira, L.A., Viegas, D.X., and Raimundo, A.M. 1997. Numerical predictions on the soil thermal effect under surface fire conditions. Int. J. Wildland Fire, 7: 51-63.

Ownbey, G.B., and Morley, T. 1991. Vascular plants of Minnesota: a checklist and atlas. University of Minnesota Press, Minneapolis, Minnesota.

Persson, H.A. 1983. The distribution and productivity of fine roots in boreal forests. Plant Soil, 71: 87-101.

Rusterholz, K. 1990. Minnesota's old growth forests. Minnesota Forests, 3: 12-16.

Singer, R. 1986. The Agaricales in modern taxonomy. 4th ed. Koeltz Scientific Books. Koenigstein, Germany.

Staddon, W.J., Duchesne, L.C., and Trevors, J.T. 1996. Conservation of forest soil microbial diversity: the impact of fire and research needs. Environ. Rev. 4: 267- 275. 59

Torres, P., and Honrubia, M. 1997. Changes and effects of a natural fire on ectomycorrhizal inoculum potential of soil in a Pinus halepensis forest. For. Ecol. Manag. 96: 189-196.

USDA Forest Service. 1998. Characteristics of the Superior National Forest landtype associations. USDA Forest Service, Superior National Forest, Duluth, Minnesota.

Villeneuve, N., Grandtner, M.M., and Fortin, J.A. 1989. Frequency and diversity of ectomycorrhizal and saprophytic macrofungi in the Laurentide Mountains of Quebec. Can. J. Bot. 67: 2616-2629.

Visser, S. 1995. Ectomycorrhizal fungal succession in jack pine stands following wildfire. New Phytol. 129: 389-401.

Wendt, K.M., and Coffin, B.A. 1988. Natural vegetation of Minnesota at the time of the public lands survey 1847-1907. Minn. Dep. Nat. Res. Biol. Rep. 1. St. Paul, Minnesota. 60

CHAPTER 4. GENERAL CONCLUSION

General discussion

After three seasons of sampling, 179 taxa were documented at the study site. Of the total taxa, 167 (63%) were true mushrooms and 12 (7%) were boletes. Identifications were made, to at least the genus level, for 156 taxa, although not all were definitive. Twenty-three taxa could not be identified (Chapter 2, Table 1). Ectomycorrhizal status was determined, based on published literature, for 149 of the 179 taxa (Chapter 2, Table 1). The _30 taxa that were not assigned an ectomycorrhizal status were the 28 unidentified species, five taxa for which the genus identification was uncertain, Naucoria sp. (#312), and Entoloma majaloides

Orton. Of the 149 taxa for which an ectomycorrhizal status was determined, 102 (68%) were ectomycorrhizal and 47 (32%) were non-mycorrhizal. Other researchers have also found that ectomycorrhizal fungi are the dominant ecological group in coniferous forests (Bills et al.

1996; Villeneuve et al. 1989). The difference between the average ectomycorrhizal species richness (unit x = 37) and the average non-mycorrhizal species richness (unit x = 17) per unit at the study site was statistically significant (P=0.004).

The dominant genera at the study site, in terms of species richness, were Russula (25 species), Cortinarius (23 species), Lactarius (11 species), and Amanita (9 species). Other authors have also reported these to be amongst the most diverse genera in conifer-dominated ecosystems (e.g., Villeneuve et al. 1989; Palmer et al. 1994; Leacock 1997). In addition to the four previously mentioned genera, Laccaria, Collybia, Hypholoma, Mycena, and

Pholiota were all dominant genera in terms of sporocarp abundance (Appendix Table 2). 61

The burned units had fewer overall species (unit x = 51) than the unburned units (unit x =75) (pooled std. dev.=3.77, P=0.006). Three-year ectomycorrhizal species richness was also lower in the burned units (unit x = 32) than in the unburned units (unit x = 45) (pooled std. dev.=3.88, P=0.036). These results indicate that the fire killed sufficient ectomycorrhizae to significantly reduce the overall species and ectomycorrhizal species richness in the burned units. Stendell et al. (1999) discovered an eight-fold reduction in ectomycorrhizal biomass present in the litter and organic horizons of burned plots compared to control plots in a ponderosa pine forest. Dahlberg et al. (2001) documented a decrease in the abundance of mycorrhizae and the number of ectomycorrhizal types after slight- and hard-bums compared to unburned Swedish boreal forest. Further results from the current study, however, demonstrate that the fire did not decimate the ectomycorrhizal fungal community in the burned units, because there was no difference in the percentage of ectomycorrhizal species occurring in the burned units (unit x = 68) compared to the unburned units (unit x =70) (pooled std. dev.=0.067, P=0.779).

Total density of sporocarps at the study site was greatest in 2000, with 14,900 sporocarps/ha. This is similar to Leacock's (1997) estimate of 14,000-16,000 sporocarps/ha for a Minnesota red pine forest. Overall sporocarp abundance did not differ between the burned (unit x =223) and unburned units (unit x =358) (pooled std. dev.=110, P=0.272),

There was also no difference in the abundance of ectomycorrhizal sporocarps in the burned

(unit x = 113) and unburned (unit x = 126) units (pooled std. dev.=33.7, P=0.719).

O'Halloran et al. (1987) found no difference between prescribed burned and unburned sites in terms of sporocarp production in a Texas pine-hardwood forest. There are many factors, e.g., precipitation, that likely influence annual and spatial fruiting patterns of basidiocarps 62

(Vogt et al. 1992). However, the results of this study indicate that fire did not negatively alter the abundance of sporocarps produced during the three-year study.

1970). In this study, they were documented exclusively in burned units.

The fruiting season started in mid-May and lasted until early October. Sporocarp production peaked in August and September (Chapter 2, Fig. 2). Leacock (1997) also found that sporocarp production in north central Minnesota was most abundant in September.

Fruiting of specific taxa was very inconsistent from year to year, and there was not much species overlap. Of the three-year taxa total, only 16 species (9%) fruited during every year of the study, while 114 species (64%) appeared in only one of the years (Appendix Table 4).

Similarly, Dahlberg et al. (1997) found that nearly 50% of the macrofungal species in a

Swedish, old-growth Picea abies forest occurred in only one year of the six-year study.

This study demonstrates that, while assumptions and speculations can be made as to the effects of fire on ectomycorrhizal fungi, it is difficult to make concrete statements. While ectomycorrhizal species richness decreased immediately following a low intensity burn, many mycorrhizae survived and produced abundances and species richness that were no different from the unburned units. Despite the fact that species with a shallow distribution of their mycelia would probably decrease after a burn (Dahlberg at al. 2001), it appears that the ectomycorrhizal fungal community, in the long-term, would not generally be adversely affected by low intensity prescribed burns. 63

Recommendations for future research

Approximately 40 of the 179 species collected during this study appear to be new records for Minnesota (D. McLaughlin, personal communication). However, this needs to be confirmed by further examination of records at the University of Minnesota Herbarium and other herbaria, a more thorough search of the literature for documentation of Minnesota fungi, and expert advice on some questionable identifications. Even if not all these species are eventually shown to be new state records, it does indicate that there is still much to be done in terms of documenting macrofungi in northeastern Minnesota and collecting baseline data. This study provides a partial picture of the macrofungal diversity of the area. Leacock

( 1997) found 148 ectomycorrhizal species in a Minnesota red pine-dominated forest. Peter et al. (2001) found 128 ectomycorrhizal species in Swiss forests dominated by P. abies. Not only did these researchers document more ectomycorrhizal species than the current study, but they did so in ecosystems with far less vegetative diversity. The current study was conducted in an area with seven dominant tree species known to form ectomycorrhizal associations.

Also, it is suggested that, when relying solely on sporocarp surveys, long-term monitoring is necessary to obtain complete characterization of species diversity (Vogt et al. 1992;

Hawksworth 2001). One would therefore expect that further sampling would reveal an even more complete picture of the macrofungal diversity of the area. 64

References

Dix, N.J., and Webster, J. 1995. Fungal ecology. Chapman & Hall, London.

Hawksworth, D.L. 2001. The magnitude of fungal diversity: the 1.5 million species estimate revisited. Mycol. Res. 105: 1422-1432.

O'Halloran, K.A., Blair, R.M., Alcaniz, R., and Hershel, F.M., Jr. 1987. Prescribed burning effects on production and nutrient composition of fleshy fungi. J. Wildl. Manag. 51: 258-262.

Petersen, P.M. 1970. Danish fireplace fungi, an ecological investigation of fungi on bums. Dan. Bot. Ark. 27: 6-97.

Smith, A.H., and Hesler, L.R. 1968. The North American spcies of Pholiota. Hafner Publishing Co., New York, New York. 65

Villeneuve, N., Grandtner, M.M., and Fortin, J.A. 1989. Frequency and diversity of ectomycorrhizal and saprophytic macrofungi in the Laurentide Mountains of Quebec. Can. J. Bot. 67: 2616-2629.

APPENDIX A.

IDENTIFICATION RESOURCES 67

Identification Resources

Resources used for identification ranged from technical keys to non-technical identification guides. While there are many books on fungi, most are mainly pictoral works which are lacking critical information necessary to make a positive identification. There are also a limited number of fungal identification resources for the northeastern United States, and many times I consulted European sources. The following is a list of references that I used, to varying degrees, to identify the fungi that were collected during the three-year study.

Also included are samples of two different data sheets on which information for each species was recorded for this study.

Barron, G. 1999. Mushrooms of northeast North America. Lone Pine Publishing, Edmonton, Alberta.

Bessette, A.E., Bessette, A.R., and Fischer, D.W. 1997. Mushrooms of northeastern North America. Syracuse University Press, Syracuse, New York.

Bessette, A.E., Miller, O.K., Jr., Bessette, A.R., and Miller, H.H. 1995. Mushrooms of North America in color: a field guide companion to seldom-illustrated fungi. Syracuse University Press, Syracuse, New York.

Bessette, A.E., Roody, W.C., and Bessette, A.R. 2000. North American boletes. Syracuse University Press, Syracuse, New York.

Breitenbach, J., and Kranzlin, R. (Editors). 1991. Fungi of Switzerland, Volume 3. Boletes and agarics, 1st part. Edition Mykologia Lucerne, Lucerne, Switzerland.

Breitenbach, J., and Kranzlin, R. (Editors). 1995. Fungi of Switzerland, Volume 4. Agarics, 2nd part. Edition Mykologia Lucerne, Lucerne, Switzerland.

Breitenbach, J., and Kranzlin, R. (Editors). 2000. Fungi of Switzerland, Volume 5. Agarics, 3rd part. Edition Mykologia Lucerne, Lucerne, Switzerland.

Courtecuisse, R. 1999. Mushrooms: a photographic guide to the mushrooms of Britain and Europe (Collins Wildlife Trust Guide). HarperCollins Publishers, London. 68

Groves, J.W. 1962. Edible and poisonous mushrooms of Canada. Canada Department of Agriculture, Ottawa, Ont.

Halling, R.E. 1983. The genus Collybia (Agaricales) in the northeastern United States and adjacent Canada. Mycologia Memoir No. 8. J. Cramer Publisher, Braunschweig, Germany.

Hansen, L., and Knudsen, H. (Editors). 1992. Nordic macromycetes. 2. Nordsvamp, Copenhagen.

Hesler, L.R., and Smith, A.H. 1963. North American species of Hyrophorus. University of Tennessee Press, Knoxville, Tennessee.

Hesler, L.R., and Smith, A.H. 1979. North American species of Lactarius. University of Michigan Press, Ann Arbor, Michigan.

Homola, R.L., and Czapowskyj, M.M. 1981. Ectomycorrhizae of Maine 2. A listing of Lactarius with the associated hosts. Maine Agricultural and Experiment Station. University of Maine, Orono, Maine. Bulletin 779.

Homola, R.L., Czapowskyj, M.M., and Blum, B.M. 1985. Ectomycorrhizae of Maine 3. A listing of Hygrophorus with the associated hosts. Maine Agricultural and Experiment Station, University of Maine, Orono, Maine. Bulletin 810.

Huffman, D.M., Tiffany, L.H., and Knaphus, G. 1989. Mushrooms and other fungi of the midcontinental United States. Iowa State University Press, Ames, Iowa.

Jenkins, D.T. 1986. Amanita of North America. Mad River Press, Eureka, California.

Kauffman, C.H. 1918. The Agaricaceae of Michigan. 1. W.H. Crawford Co., Lansing, Michigan.

Kibby, G. 1992. Mushrooms and other fungi. Smithmark Publishers Inc., New York, New York.

Kibby, G., and Fatto, R. 1990. Keys to the species of Russula in northeastern North America. 3rd ed. Kibby-Fatto Enterprises, Somerville, New Jersey.

Lress0e, T., and Lincoff, G. 1998. Mushrooms. Dorling Kindersley Ltd., London.

Lange, M., and Hora, F.B. 1963. A guide to mushrooms and toadstools. E.P. Dutton & Co., Inc., New York, New York.

Largent, D.L. 1977. How to identify mushrooms to genus I: macroscopic features. Mad River Press, Eureka, California. 69

Largent, D.L., and Thiers, H.D. 1977(?). How to identify mushrooms to genus II: field identification of genera. Mad River Press, Eureka, California.

Lincoff, G.H. 1998. National Audubon Society field guide to North American mushrooms. Alfred A. Knopf, Inc., New York, New York.

Miller, O.K., Jr. 1978. Mushrooms of North America. E.P. Dutton & Co., Inc., New York, New York.

Moser, M. 1983. Keys to agarics and boleti. Roger Phillips, London.

Mueller, G.M. 1992. Systematics of Laccaria (Agaricales) in the continental United States and Canada, with discussions on extralimital taxa and descriptions of extant types. Fieldiana, Botany, New Series No. 30.

Neville, J. 1996. How identification of "little brown mushroom" (LBM) genera may not be as hard as you think. Mcllvainea, 12: 33-63.

Orton, P.D., and Watling, R. 1979. British fungus flora, agarics and boleti part 2: Coprinaceae: Coprinus. Her Majesty's Stationery Office, Edinburgh, Scotland.

Pacioni, G. 1981. Simon and Schuster's guide to mushrooms. Simon & Schuster, Inc. New York, New York.

Phillips, R. 1981. Mushrooms and other fungi of Great Britain and Europe. Pan Books, Ltd., London.

Phillips, R. 1991. Mushrooms of North America. Little, Brown & Co., Boston, Mass.

Smith, A.H., and Hesler, L.R. 1968. The North American species of Pho/iota. Hafner Publishing Co., New York, New York.

Smith, A.H., and Weber, N.S. 1996. The mushrooms hunter's field guide. University of Michigan Press, Ann Arbor, Michigan.

Smith, A.H., Smith, H.V., and Weber, N.S. 1979. How to know the gilled mushrooms. Wm. C. Brown Co., Dubuque, Iowa.

Stuntz, D .E. 1977. How to identify mushrooms to genus IV: keys to families and genera. Mad River Press, Eureka, California.

Weaver, M.G., and McLaughlin, D.J. 1980. Mushroom flora of Minnesota - a contribution. Bell Museum of Natural History, University of Minnesota, Minneapolis. Occasional Papers: 16. 70

Genus Species ______Collection# ______

Plot# Date Tally ______# Col. ______Plot# Date Tally #Col. ______Plot# Date Tally #Col. ______Plot# Date Tally #Col. ______Plot# Date Tally #Col. ______Plot# Date Tally # Col. ______

I CAP/PILEUS: III STIPE cont. 0 Color:______0 Terete 0 Convex D Compressed 0 Conical 0 Campanulate D Equal 0 Broadly parabolic D Clavate 0 Narrow parabolic D Bulbuous 0 Umbonate D Taper-base to apex 0 Broadly umbonate D Taper-apex to base 0 Shallow depressed . 0 Uplifted D Volva: 0 Plane 0 Annulus:

0 Smooth D Hollow 0 Waxy D Stuffed 0 Velvety D Solid 0 Fibrous 0 Loose scales D Reticulate 0 Seal es (in temal) / Cracks D Fibrous 0 Striate D Glandular-dotted 0 Pleated IV MARGIN: 0 Dry D lruolled 0 Gelatinous D Scalloped 0 Viscid D Undulating

0 Flesh color/bruising: V GROWfH HABIT: D Solitary 0 Other: D Scattered 0 Gregarious II GILLS/PORES: 0 Caespitose 0 Color: 0 Free VI MEASUREMENTS: 0 Attached Cap Size: 0 Sinuate 0 Subdecurrent Stipe Lenth: 0 Decurrent 0 Adnexed Stipe Width: 0 Adnate Tube Length & per/ mm: 0 Distant 0 Close Spore Size: (40X) 0 Crowded Spore Wall: 0 Equal 0 Unequal 0 Forked D Oil Drops: D Cystidia: III STIPE: D Spaerocysts 0 Color: 0 Central Spore Print: 0 Lateral 71

Genus Species Collection #

. Plot# Date Tally # Col. Plot# Date Tally # Col. ~ Plot# Date Tally # Col. Plot# Date Tally #Col. Plot# Date Tally # Col. Plot# Date Tally #Col. Plot# Date Tally #Col. Plot# Date Tally #Col.

Field Notes:

I CAP/PILEUS: III STIP'E: 0 Color: 0 Color: 0 Shape: 0 Shape (x-sec.) 0 Shape (long.) 0 Surface: 0 Color changes: 0 Consistency: 0 Flesh: 0 Margin: 0 Volva: Shape (x-section) Type Shape (surface) Color Surface 0 Annulus/Veil 0 Surface: Type Shininess Color Wetness Texture IV GROWTH HABIT: 0 0 Cuticle Peel: Solitary 0 Scattered 0 Flesh 0 Gregarious Color &/ or changes 0 Caespitose V MEASUREMENTS: Thickness Cap Size: Taste/Odor Stipe Length: Stipe Width: 0 Other: Tube Length & per/mm:

II GILLS/PORES: Spore Size: (40X) 0 Color: 0 Attachment: Spore Wall Texture: To stipe Spore Characteristics: Margin to stipe 0 Spacing: Spore Print 0 Margin: 0 Taste: 0 Separation from cap: 72

APPENDIXB.

ADDITIONAL TABLES 73

Table !. -Number of species d-ocumented Table 2. Number of sporocarps coilected - per genus during the three-year study, for each genus during the three-year study, and the relative frequency of each genus. and the relative frequency of the genus. Genus Species Relative Genus Sporocarp Relative Richness Frequency Richness Frequency (%) ( %)

Russula 25 17 Collybia 539 13 Cortinarius 23 15 Cortinarius 469 12 Lactarius 11 7 Lactarius 329 10 Amanita 9 6 Laccaria 305 8 Mycena 8 5 Russula 292 7 Hygrocybe 7 5 Hypholoma 288 7 Suillus 7 5 Mycena 281 7 Tricholoma 7 5 Pho/iota 266 7 Collybia 6 4 Hygrocybe 199 5 Inocybe 6 4 Armillaria 171 4 Leptonia 6 4 Myxopmphalia 132 3 Clitocybe 4 3 Marasmius 129 3 Hygrophorus 3 2 Suillus 113 3 Boletus 2 1 Chrysomphalina 98 2 Hebeloma 2 1 Amanita 59 1 Hypholoma 2 1 Inocybe 57 1 Laccaria 2 1 Tricholoma 53 1 Marasmius 2 1 Leptonia 41 1 Myxopmphalia 2 1 Hygrophorus 37 1 Nolanea 2 1 Coprinellus 30 1 Phaeocollybia 2 1 Clitocybe 22 0.5 Armillaria 1 0.8 Phaeocollybia 21 0.5 Chroogomphus 1 0.8 Rozites 12 0.3 Chrysomphalina 1 0.8 Hebeloma 11 0.3 Coprinellus 1 0.8 Entoloma 10 0.3 Entoloma 1 0.8 Xeromphalina 10 0.3 Leccinum 1 0.8 Tylopilus 5 0.1 Naucoria 1 0.8 Boletus 4 0.1 Pho/iota 1 0.8 Nolanea 3 0.1 Rozites 1 0.8 Tricholomopsis 3 0.1 Tricholomopsis 1 0.8 Chroogomphus 2 0.05 Tylopilus 1 0.8 Leccinum 1 0.02 Xeromphalina 1 0.8 Naucoria 1 0.02 74

Table 3. Distribution of epigeous macrofungi amongst burned units (Units 1-3) and unburned units (Units 4-5) during the three-year study. Collection Species Unit 1 Unit 2 Unit 3 Unit 4 Unit 5 Number Boletales 124/149 Boletus subtomentosus L. X X 313 Boletus sp. X 10 Chroogomphus corallinus O.K. Miller & Wat!. X 150 Leccinum aurantiacum (Bull. ex St. Am.) S.F. Gray X 141/175/184 Suillus granulatus (L.:Fr.) Roussel X X X M/N/132/135 Suillus pictus (Peck) A.H. Sm. & Theirs X X X X 128 Suillus placidus (Bonord.) Sing. X 158 Suillus subalutaceous (A.H. Sm. & Thiers) X X A.H. Sm. & Thiers K Suillus sp. X z Suillus sp. X 37 Suillus sp. X y Suillus sp. (?) X X X X 129/311 Tylopilus felleus (Bull.:Fr.) Karst. X X X X

Agaricales 308 Amanita ceciliae (Berk. & Br.) Bas X 117 Amanita flavoconia Atk. X 154/185 Amanitafulva (Schaeff.) Pers. X X X H Amanitafulva (?) (Schaeff.) Pers. X 203 Amanita porphyria (Alb. & Schw.:Fr.) Mlady X 134 Amanita vaginata (Bull. ex Fr.) Que!. X F Amanita virosa (Fr.) G. Berto!. X X X X R Amanita sp. X 206 Amanita sp. X L/T/188 Armillaria mellea complex (Vahl:Fr.) Kumm. X X X X X 14 Chrysomphalina sp. X 182/331 Clitocybe clavipes (Pers.:Fr.) Kumm. X X X 208 Clitocybe sinopicoides Peck X 148 Clitocybe vilescens (?) Peck X 213 Clitocybe sp. X 8/204 Collybia butyracea (Bull.:Fr.) Kumm. X X X 6/119 Collybia confluens (Pers.:Fr.) Kumm. X X X 104/115 Collybia dryophila (Bull.:Fr.) Kumm. X X X X X 305/306 108/300/302 Collybia luteifolia Gill. X X X 169 Collybia maculata (Alb. & Schw.:Fr.) Kumm. X 160 Collybia proxilla (Fr.) Gill. X 33/101 Coprinellus angulatus Peck X 170 Entoloma majaloides Orton X 155 Hygrocybe acutoconica var. microspora (?) X Hesler & A.H. Sm. 315 Hygrocybe conica (Scop.:Fr.) Kumm. var. conica X 123/140 Hygrocybe laeta (Pers.:Fr.) Kumm. X X X X 75

Table 3. Continued Collection Species Unit 1 Unit 2 Unit 3 Unit4 Unit 5 Number 138 Hygrocybe marginata Peck var. marginata X X 118 Hygrocybe miniata (Fr.) Kumm. X X X 194 Hygrocybe miniata (Berk. & Br.) E. Arnolds var. mollis (?) X X X 162 Hygrocybe mini at a (?) (Fr.) Kumm. X G/21/172 Hygrophorus angustifolius (Murr.) Hesler & A.H. Sm. X X 334 Hygrophorus camarophyllus (Alb. & Schw.:Fr.) X X Dumee, Grandjean & Maire 40 Hygrophorus russula (Scop.:Fr.) Que!. X 195 Hygrophorus sp. (?) X 338 Hygrophorus sp. (?) X 2 Hypholoma fasciculare (Huds. :Fr.) Kumm. X 100 Hypholomafasciculare (?) (Huds.:Fr.) Kumm. X 159 Laccaria bicolor (Maire) Orton X 4/122/133 Laccaria laccata (Scop.:Fr.) Berk. & Br. X X X X X 147 144 Leptoniaformosus (Fr.) Que!. X 310 Leptoniafulvus (?) (Orton) Mos. X 320 Leptonia sericellum (Bull. ex Fr.) Kumm. X 43 Leptonia sp. X 145 Leptonia sp. X X 199 Leptonia sp. X 109 Marasmius androsaceus (L.:Fr.) Fr. X 111 Marasmius sp. X 173 Mycena epipterygia var. pelliculosa (?) (Fr.) Maas G. X 210 Mycena griseoviridis A.H. Sm. X 36 Mycena griseoviridis (?) A.H. Sm. X 103 Mycena sanguinolenta (Alb. & Schw.:Fr.) Kumm X X X X 102 Mycena sp. X X X X X 179 Mycena sp. X 307 Mycena sp. X X 319 Mycena sp. X 177/303 Myxomphalia maura (Fr.) Hora X 34 Myxomphalia maura (?) (Fr.) Hora X 312 Naucoria sp. X 143 Nolanea nitens (?) (Ve!.) K. & R. X 116 Nolanea sp. X 107/301/304 Pho/iota highlandensis (Peck) A.H. Sm. & Hessler X X 207 Tricholoma apium (?) Schaff. X X 187 Tricholoma columbetta (Fr.) Kumm. X X 209 Tricholomaflavovirens (Pers.:Fr.) Lundell X 186 Tricholoma imbricatum (?) (Fr.:Fr.) Kumm. X 330 Tricholoma magnivelare (Peck) Redhead X 205 Tricholoma saponaceum (Fr.:Fr.) Kumm. X 45 Tricholoma sejunctum (Sow.:Fr.) Que!. X X 120 Tricholomopsis rutilans (Schaeff.:Fr.) Sing. X 76

Table 3. Continued Collection Species Unit 1 Unit 2 Unit 3 Unit4 Unit 5 Number 114 Xeromphalina cauticinalis Kiihn. & Maire X X X X

Cortinariales 7b/163/l 97 Cortinarius alboviolaceus (Pers.:Fr.) Fr. X X X 0/168 Cortinarius armillatus (Fr. :Fr.) Fr. X 29/46 Cortinarius bolaris (Pers.:Fr.) Fr. X X X 178 Cortinarius cinnamomeus (L.:Fr.) Fr. X 41/181 Cortinarius leucopus (?) (Bull. ex Fr.) Fr. X X X X X 183 Cortinarius lividoochraceus (?) (Berk.) Berk. X 23a Cortinarius multiformis var. coniferarum Mos. X 7a Cortinarius obliquus Peck X X X X 151/211 Cortinarius obtusus (?) (Fr.) Fr. X X X 333 Cortinarius rubricosus (?) Fr. X 12 Cortinarius semisanguineus (Fr.) Gill. X X 176 Cortinarius sphaerosporus Peck X 142/164 Cortinarius subtorvus (?) Lamoure X X 161 Cortinarius tenebricus (?) Favre X X X X 202/322 Cortinarius trivia/is Lange X 1/27 Cortinarius sp. X X X X 15 Cortinarius sp. X X 23b Cortinarius sp. X 152 Cortinarius sp. X 193 Cortinarius sp. X 196 Cortinarius sp. X 317 Cortinarius sp. X 332 Cortinarius sp. X 214 Cortinarius sp. (?) X X 318 Cortinarius sp. (?) X 192 Hebeloma crustuliniforme (Bull.) Que!. X X 136 Hebeloma sinapizans (?) (Paulet) Gill. X 167 Hebeloma sp. (?) X 139 Jnocybe lacera (Fr.) Kumm. X 121 Inocybe leptophylla Atk. X X 309 Inocybe proximella (?) Karst. X 24/335 Jnocybe sp. X 25 Jnocybe sp. X X X 214 Jnocybe sp. X 9 Phaeocollybia christinae (Fr.) Heim X 11 Phaeocollybiajennyae (Karst.) Heim X 191 Rozites caperatus (Pers.:Fr.) Karst. X X

Russulales 131/174 Lactarius a/finis var. a/finis Peck X X 22/156/165 Lactarius camphoratus (Bull.:Fr.) Fr. X X X X 323 Lactarius deceptivus Peck X X 329 Lactarius hibbardae var. hibbardae Peck X X 77

Table 3. Continued Collection Species Unit 1 Unit 2 Unit 3 Unit 4 Unit 5 Number 106 Lactarius lignyotus Fr. X 324 Lactarius sordidus Peck X 190 Lactarius subvellereus var. subvellereus Peck X 189 Lactarius torminosus (Schaeff.:Fr.) Pers. X 166 Lactarius uvidus (Fr.) Fr. var. uvidus X 5/180 Lactarius vinaceorufescens A.H. Sm. X X X X X AA/E/26 Lactarius sp. X X X X X 126 Russula bicolor (?) Burl. X 125 Russula brevipies Peck X X 130 Russula brunneola Burl. X 112/198 Russula cyanoxantha (Schaeff.) Fr. f. cyanoxantha X X 31 Russula decolorans (?) (Fr.) Fr. X X 137 Russula elaeodes (Bres.) Bon X 327 Russula faginea (?) Romagn. X 13 Russula fragilis (Pers.: Fr.) Fr. X X X X 3/146 Russulafragrantissima Romagn. X X X 326 Russula incarnaticeps (?) Murrill X 328 Russula paludosa Britz. X 157 Russula paludosa (?) Britz. X 337 Russula paludosa (?) Britz. X X X 336 Russula sanguinea (Bull.) Fr. X 19 Russula sanguinea (?) (Bull.) Fr. X X X X J Russula sp. X 38 Russula sp. X 171 Russula sp. X 200 Russula sp. X 314 Russula sp. X X 325 Russula sp. X 339 Russula sp. X 340 Russula sp. X 341 Russula sp. X 342 Russula sp. X

A/B Unknown X X C Unknown - Cortinarius sp. (?) X p Unknown X Q Unknown X s Unknown X u Unknown X V Unknown X X w Unknown X X Unknown X 16 Unknown X X 17 Unknown X 18 Unknown X 28 Unknown X 78

Table 3. Continued Collection Species Unit 1 Unit 2 Unit 3 Unit 4 Unit 5 Number 32 Unknown X 35 Unknown X 39 Unknown X 42 Unknown X 110 Unknown X 113 Unknown X 153 Unknown X 201 Unknown X 215 Unknown X 321 Unknown X

UNIT TOTALS 52 54 46 77 73 79

Table 4. Numbers of sporocarps per species collected each year during the three-year study. Collection Species 1999 2000 2001 Total Number Sporocarps Boletales 124/149 Boletus subtomentosus L. 2 2 313 Boletus sp. 2 2 10 Chroogomphus corallinus O.K. Miller & Watl. 2 2 150 Leccinum aurantiacum (Bull. ex St. Am.) S.F. Gray 1 141/175/184 Suillus granulatus (L.:Fr.) Roussel 5 7 12 M/N/132/135 Suillus pictus (Peck) A.H. Sm. & Theirs 15 7 22 128 Suillus placidus (Bonord.) Sing. 5 5 158 Suillus subalutaceous (A.H. Sm. & Thiers) 4 4 A.H. Sm. & Thiers K Suillus sp. 67 67 z Suillus sp. 1 1 37 Suillus sp. 2 2 y Suillus sp. (?) 24 24 129/311 Tylopilus felleus (Bull.:Fr.) Karst. 4 5

Agaricales 308 Amanita ceciliae (Berk. & Br.) Bas 117 A manila flavoconia Atk. 4 10 14 154/185 Amanitafulva (Schaeff.) Pers. 11 9 20 H Amanita fulva (?) (Schaeff.) Pers. 9 9 203 Amanita porphyria (Alb. & Schw.:Fr.) Mlady 1 134 Amanita vaginata (Bull. ex Fr.) Quel. 2 F Amanita virosa (Fr.) G. Bertot. 4 6 10 R Amanita sp. 206 Amanita sp. l L/T/188 Armillaria me/lea complex (Vahl:Fr.) Kumm. 140 9 22 171 14 Chrysomphalina sp. 50 48 98 182/331 Clitocybe clavipes (Pers.:Fr.) Kumm. 2 5 7 208 Clitocybe sinopicoides Peck 2 2 148 Clitocybe vilescens (?) Peck 1 1 12 213 Clitocybe sp. 1 8/204 Collybia butyracea (Bull.:Fr.) Kumm. 14 14 2 30 6/119 Collybia confluens (Pers.:Fr.) Kumm. 15 240 24 279 104/115/305/306 Collybia dryophila (Bull.:Fr.) Kumm. 59 151 210 108/300/302 Collybia luteifolia Gill. 3 10 13 169 Collybia maculata (Alb. & Schw.:Fr.) Kumm. 6 6 160 Collybia proxilla (Fr.) Gill. 1 33/101 Coprinellus angulatus Peck 15 14 30 170 Entoloma majaloides Orton 10 10 155 Hygrocybe acutoconica var. microspora (?) 7 7 Hesler & A.H. Sm. 315 Hygrocybe conica (Scop.:Fr.) Kumm. var. conica 123/140 Hygrocybe laeta (Pers.:Fr.) Kumm. 69 29 98 138 Hygrocybe marginata Peck var. marginata 5 7 12 80

Table 4. Continued Collection Species 1999 2000 2001 Total Number Sporocarps 118 Hygrocybe miniata (Fr.) Kumm. 56 9 65 194 Hygrocybe miniata (Berk. & Br.) E. Arnolds var. mollis (?) 10 2 12 162 Hygrocybe miniata (?) (Fr.) Kumm. 4 4 G/21/172 Hygrophorus angustifolius (Murr.) Hesler & A.H. Sm. 7 12 1 1 30 334 Hygrophorus camarophyllus (Alb. & Schw.:Fr.) 4 4 Dumee, Grandjean & Maire 40 Hygrophorus russula (Scop.:Fr.) Quel. 2 3 195 Hygrophorus sp. (?) 2 2 338 Hygrophorus sp. (?) 3 3 2 Hypholoma fasciculare (Huds. :Fr.) Kumm. 58 150 66 274 100 Hypholomafasciculare (?) (Huds.:Fr.) Kumm. 14 14 159 Laccaria bi color (Maire) Orton 1 2 4/122/133/147 Laccaria laccata (Scop.:Fr.) Berk. & Br. 57 235 11 303 144 Leptoniaformosus (Fr.) Quel. 1 1 310 Leptonia fulvus (?) (Orton) Mos. 6 6 320 Leptonia sericellum (Bull. ex Fr.) Kumm. 2 2 43 Leptonia sp. 19 19 145 Leptonia sp. 10 10 199 Leptonia sp. 3 3 109 Marasmius androsaceus (L.:Fr.) Fr. X (hundreds) 111 Marasmius sp. 54 75 129 173 Mycena epipterygia var. pelliculosa (?) (Fr.) Maas G. 4 4 210 Mycena griseoviridis A.H. Sm. 1 36 Mycena griseoviridis (?) A.H. Sm. 2 2 103 Mycena sanguinolenta (Alb. & Schw.:Fr.) Kumm 127 16 143 102 Mycena sp. 112 112 179 Mycena sp. 17 17 307 Mycena sp. 4 4 319 Mycena sp. 1 177/303 Myxomphalia maura (Fr.) Hora 124 2 126 34 Myxomphalia maura (?) (Fr.) Hora 6 6 312 Naucoria sp. 143 Nolanea nitens (?) (Vel.) K. & R. 2 2 116 Nolanea sp. 1 1 107/301/304 Pho/iota highlandensis (Peck) A.H. Sm. & Hessler 190 76 266 207 Tricholoma apium (?) Schaff. 2 187 Tricholoma columbetta (Fr.) Kumm. 4 18 22 209 Tricholomaflavovirens (Pers.:Fr.) Lundell 186 Tricholoma imbricatum (?) (Fr.:Fr.) Kumm. 330 Tricholoma magnivelare (Peck) Redhead 205 Tricholoma saponaceum (Fr. :Fr.) Kumm. I 2 3 45 Tricholoma sejunctum (Sow.:Fr.) Quel. 9 9 5 23 120 Tricholomopsis rutilans (Schaeff.:Fr.) Sing. 3 3 114 Xeromphalina cauticinalis Kiihn. & Maire 10 10 81

Table 4. Continued Collection Species 1999 2000 2001 Total Number Sporocarps Cortinariales 7b/l 63/l 97 Cortinarius alboviolaceus (Pers.:Fr.) Fr. 14 32 15 61 D/168 Cortinarius armillatus (Fr.:Fr.) Fr. 11 40 6 57 29/46 Cortinarius bolaris (Pers.:Fr.) Fr. 12 18 30 178 Cortinarius cinnamomeus (L. :Fr.) Fr. 8 6 14 41/181 Cortinarius leucopus (?) (Bull. ex Fr.) Fr. 25 56 18 99 183 Cortinarius lividoochraceus (?) (Berk.) Berk. 2 2 23a Cortinarius multiformis var. coniferarum Mos. 2 I 3 7a Cortinarius obliquus Peck 12 5 6 23 151/211 Cortinarius obtusus (?) (Fr.) Fr. 25 25 333 Cortinarius rubricosus (?) Fr. 4 4 12 Cortinarius semisanguineus (Fr.) Gill. 27 24 51 176 Cortinarius sphaerosporus Peck I 1 142/164 Cortinarius subtorvus (?) Lamoure IO 10 161 Cortinarius tenebricus (?) Favre 18 18 202/322 Cortinarius trivia/is Lange 2 1/27 Cortinarius sp. 33 34 15 Cortinarius sp. 5 5 23b Cortinarius sp. 2 2 152 Cortinarius sp. 193 Cortinarius sp. I 196 Cortinarius sp. 16 16 317 Cortinarius sp. 7 7 332 Cortinarius sp. 3 3 214 Cortinarius sp. (?) 14 14 318 Cortinarius sp. (?) 192 Hebeloma crustuliniforme (Bull.) Quel. 10 10 136 Hebeloma sinapizans (?) (Paulet) Gill. I 167 Hebeloma sp. (?) 2 2 139 lnocybe lacera (Fr.) Kumm. 1 1 I 12 121 !nocybe leptophylla Atk. 5 3 8 309 Jnocybe proximella (?) Karst. 7 7 24/335 lnocybe sp. 2 10 12 25 lnocybe sp. 15 2 17 214 Jnocybe sp. 9 Phaeocollybia christinae (Fr.) Heim 1 11 Phaeocollybiajennyae (Karst.) Heim 11 9 20 191 Rozites caperatus (Pers.:Fr.) Karst. 4 8 12

Russulales 131/174 Lactarius affinis var. affinis Peck 6 7 22/156/165 Lactarius camphoratus (Bull.:Fr.) Fr. 31 26 58 323 Lactarius deceptivus Peck 9 9 329 Lactarius hibbardae var. hibbardae Peck 2 2 106 Lactarius lignyotus Fr. 2 2 324 Lactarius sordidus Peck 2 2 82

Table 4. Continued Collection Species 1999 2000 2001 Total Number Sporocarps 190 Lactarius subvellereus var. subvellereus Peck 10 2 12 189 Lactarius torminosus (Schaeff. :Fr.) Pers. 1 166 Lactarius uvidus (Fr.) Fr. var. uvidus 1 1 5/180 Lactarius vinaceorufescens A.H. Sm. 14 70 75 159 AA/E/26 Lactarius sp. 59 17 76 126 Russula bicolor (?) Burl. 5 5 125 Russula brevipies Peck 9 4 13 " 130 Russula brunneola Burl. 2 2 4 112/198 Russula cyanoxantha (Schaeff.) Fr. f. cyanoxantha 4 5 31 Russula decolorans (?) (Fr.) Fr. 2 3 137 Russula elaeodes (Bres.) Bon 2 327 Russula faginea (?) Romagn. 1 1 13 Russulafragilis (Pers.:Fr.) Fr. 22 41 96 159 3/146 Russula fragrantissima Romagn. 10 7 18 326 Russula incarnaticeps (?) Murrill 328 Russula paludosa Britz. 4 4 157 Russula paludosa (?) Britz. 2 3 337 Russula paludosa (?) Britz. 7 7 336 Russula sanguinea (Bull.) Fr. 19 Russula sanguinea (?) (Bull.) Fr. 38 38 J Russula sp. 38 Russula sp. 2 2 4 171 Russula sp. 3 3 200 Russula sp. 2 2 314 Russula sp. 4 5 9 325 Russula sp. 339 Russula sp. 340 Russula sp. 341 Russula sp. 2 2 342 Russula sp.

A/B Unknown 3 3 C Unknown - Cortinarius sp. (?) p Unknown Q Unknown s Unknown 39 39 u Unknown 8 8 V Unknown 8 8 w Unknown X Unknown 2 2 16 Unknown 4 5 17 Unknown 2 2 18 Unknown 1 1 28 Unknown 3 3 32 Unknown 3 3 35 Unknown 16 16 83

Table 4. Continued Collection Species 1999 2000 2001 Total Number Sporocarps 39 Unknown 1 1 42 Unknown 3 3 110 Unknown 6 6 113 Unknown 2 2 153 Unknown 1 1 201 Unknown 5 5 215 Unknown 2 2 321 Unknown 2 2

SPOROCARPS: ANNUAL AND 3-YEAR TOTALS 936 2240 979 4155 SPECIES: ANNUAL AND 3-YEAR TOTALS 62 113 85 179 84

Table 5. Monthly fruiting patterns of all epigeous macrofungal species collected during the three-year study. Collection Species May June July Aug. Sept. Oct. Number Boletales 124/149 Boletus subtomentosus L. X X 313 Boletus sp. X 10 Chroogomphus corallinus O.K. Miller & Wat!. X 150 Leccinum aurantiacum (Bull. ex St. Am.) S.F. Gray X 141/175/184 Suillus granulatus (L. :Fr.) Roussel X X M/N/ 132/135 Suillus pictus (Peck) A.H. Sm. & Theirs X 128 Suillus placidus (Bonord.) Sing. X X 158 Suillus subalutaceous (A.H. Sm. & Thiers) X A.H. Sm. & Thiers K Suillus sp. X z Suillus sp. X 37 Suillus sp. X y Suillus sp. (?) X 129/311 Tylopilus felleus (Bull.:Fr.) Karst. X X

Agaricales 308 Amanita ceciliae (Berk. & Br.) Bas X 117 Amanita flavoconia Atk. X X X 154/185 Amanitafulva (Schaeff.) Pers. X X H Amanitafulva (?) (Schaeff.) Pers. X 203 Amanita porphyria (Alb. & Schw.:Fr.) Mlady X 134 Amanita vaginata (Bull. ex Fr.) Que!. X X F Amanita virosa (Fr.) G. Berto!. X X R Amanita sp. X 206 Amanita sp. X L/T/188 Armillaria mellea complex (Vahl:Fr.) Kumm. X 14 Chrysomphalina sp. X X 182/331 Clitocybe clavipes (Pers.:Fr.) Kumm. X 208 Clitocybe sinopicoides Peck X 148 Clitocybe vilescens (?) Peck X X 213 Clitocybe sp. X 8/204 Collybia butyracea (Bull.:Fr.) Kumm. X 6/119 Collybia confluens (Pers.:Fr.) Kumm. X X X 104/115 Collybia dryophila (Bull.:Fr.) Kumm. X X X X 305/306 108/300/302 Collybia luteifolia Gill. X X X 169 Collybia maculata (Alb. & Schw.:Fr.) Kumm. X 160 Collybia proxilla (Fr.) Gill. X 33/101 Coprinellus angulatus Peck X X X 170 Entoloma majaloides Orton X 155 Hygrocybe acutoconica var. microspora (?) X Hesler & A.H. Sm. 315 Hygrocybe conica (Scop.:Fr.) Kumm. var. conica X 123/140 Hygrocybe laeta (Pers.:Fr.) Kumm. X X X 85

Table 5. Continued Collection Species May June July Aug. Sept. Oct. Number 138 Hygrocybe marginata Peck var. marginata X 118 Hygrocybe miniata (Fr.) Kumm. X X X 194 Hygrocybe miniata (Berk. & Br.) E. Arnolds X var. mollis (?) 162 Hygrocybe miniata (?) (Fr.) Kumm. X G/21/172 Hygrophorus angustifolius (Murr.) Hesler & A.H. Sm. X X 334 Hygrophorus camarophyllus (Alb. & Schw.:Fr.) X X Dumee, Grandjean & Maire 40 Hygrophorus russula (Scop.:Fr.) Quel. X 195 Hygrophorus sp. (?) X 338 Hygrophorus sp. (?) X 2 Hypholoma fasciculare (Huds. :Fr.) Kumm. X X 100 Hypholomafasciculare (?) (Huds.:Fr.) Kumm. X 159 Laccaria bicolor (Maire) Orton X X 4/122/133 Laccaria laccata (Scop.:Fr.) Berk. & Br. X X X 147 144 Leptoniaformosus (Fr.) Quel. X 310 Leptoniafulvus (?) (Orton) Mos. X 320 Leptonia sericellum (Bull. ex Fr.) Kumm. X 43 Leptonia sp. X 145 Leptonia sp. X X 199 Leptonia sp. X 109 Marasmius androsaceus (L.:Fr.) Fr. X 111 Marasmius sp. X 173 Mycena epipterygia var. pelliculosa (?) (Fr.) Maas G. X 210 Mycena griseoviridis A.H. Sm. X 36 Mycena griseoviridis (?) A.H. Sm. X 103 Mycena sanguinolenta (Alb. & Schw.: Fr.) Kumm X X 102 Mycena sp. X X 179 Mycena sp. X 307 Mycena sp. X 319 Mycena sp. X 177/303 Myxomphalia maura (Fr.) Hora X X X 34 Myxomphalia maura (?) (Fr.) Hora X 312 Naucoria sp. X 143 Nolanea nitens (?) (Vel.) K. & R. X 116 Nolanea sp. X 107/301/304 Pho/iota highlandensis (Peck) A.H. Sm. & Hessler X X X X X X 207 Tricholoma apium (?) Schaff. X 187 Tricholoma columbetta (Fr.) Kumm. X 209 Tricholomaflavovirens (Pers.:Fr.) Lundell X 186 Tricholoma imbricatum (?) (Fr.:Fr.) Kumm. X 330 Tricholoma magnivelare (Peck) Redhead X 205 Tricholoma saponaceum (Fr.:Fr.) Kumm. X 45 Tricholoma sejunctum (Sow.:Fr.) Quel. X 120 Tticholomopsis rutilans (Schaeff.:Fr.) Sing. X 86

Table 5. Continued Collection Species May June July Aug. Sept. Oct. Number 114 Xeromphalina cauticinalis Ktihn. & Maire X X

Cortinariales 7b/163/l 97 Cortinarius alboviolaceus (Pers.:Fr.) Fr. X X X D/168 Cortinarius armillatus (Fr. :Fr.) Fr. X X 29/46 Cortinarius bolaris (Pers.:Fr.) Fr. X X 178 Cortinarius cinnamomeus (L.:Fr.) Fr. X X 41/181 Cortinarius leucopus (?) (Bull. ex Fr.) Fr. X X 183 Cortinarius lividoochraceus (?) (Berk.) Berk. X 23a Cortinarius multiformis var. coniferarum Mos. X 7a Cortinarius obliquus Peck X X 151/211 Cortinarius obtusus (?) (Fr.) Fr. X X X 333 Cortinarius rubricosus (?) Fr. X 12 Cortinarius semisanguineus (Fr.) Gill. X X 176 Cortinarius sphaerosporus Peck X 142/164 Cortinarius subtorvus (?) Lamoure X X 161 Cortinarius tenebricus (?) Favre X X 202/322 Cortinarius trivia/is Lange X 1/27 Cortinarius sp. X 15 Cortinarius sp. X 23b Cortinarius sp. X 152 Cortinarius sp. X 193 Cortinarius sp. X 196 Cortinarius sp. X 317 Cortinarius sp. X 332 Cortinarius sp. X 214 Cortinarius sp. (?) X 318 Cortinarius sp. (?) X 192 Hebeloma crustuliniforme (Bull.) Quel. X 136 Hebeloma sinapizans (?) (Paulet) Gill. X 167 Hebeloma sp. (?) X 139 Jnocybe lacera (Fr.) Kumm. X X 121 Jnocybe leptophylla Atk. X X 309 Jnocybe proximella (?) Karst. X X 24/335 Jnocybe sp. X 25 Jnocybe sp. X X 214 Jnocybe sp. X 9 Phaeocollybia christinae (Fr.) Heim X 11 Phaeocollybiajennyae (Karst.) Heim X 191 Rozites caperatus (Pers.:Fr.) Karst. X X

Russulales 131/174 Lactarius affinis var. affinis Peck X X 22/156/165 Lactarius camphoratus (Bull.:Fr.) Fr. X X 323 Lactarius deceptivus Peck X 329 Lactarius hibbardae var. hibbardae Peck X X 87

Table 5. Continued Collection Species May June July Aug. Sept. Oct. Number 106 Lactarius lignyotus Fr. X 324 Lactarius sordidus Peck X 190 Lactarius subvellereus var. subvellereus Peck X 189 Lactarius torminosus (Schaeff.:Fr.) Pers. X X 166 Lactarius uvidus (Fr.) Fr. var. uvidus X 5/180 Lactarius vinaceorufescens A.H. Sm. X X X AA/E/26 Lactarius sp. X 126 Russula bicolor (?) Burl. X 125 Russula brevipies Peck X X 130 Russula brunneola Burl. X X 112/198 Russula cyanoxantha (Schaeff.) Fr. f. cyanoxantha X X X 31 Russula decolorans (?) (Fr.) Fr. X X 137 Russula elaeodes (Bres.) Bon X X 327 Russula faginea (?) Romagn. X 13 Russulafragilis (Pers.:Fr.) Fr. X X 3/146 Russulafragrantissima Romagn. X X 326 Russula incarnaticeps (?) Murrill X 328 Russula paludosa Britz. X 157 Russula paludosa (?) Britz. X X 337 Russula paludosa (?) Britz. X 336 Russula sanguinea (Bull.) Fr. X 19 Russula sanguinea (?) (Bull.) Fr. X J Russula sp. X 38 Russula sp. X X 171 Russula sp. X 200 Russula sp. X 314 Russula sp. X X X 325 Russula sp. X 339 Russula sp. X 340 Russula sp. X 341 Russula sp. X 342 Russula sp. X

A/B Unknown X C Unknown - Cortinarius sp. (?) X p Unknown X Q Unknown X s Unknown X u Unknown X V Unknown X w Unknown X X Unknown X 16 Unknown X 17 Unknown X 18 Unknown X 28 Unknown X 88

Table 5. Continued Collection Species May June July Aug. Sept. Oct. Number 32 Unknown X 35 Unknown X 39 Unknown X 42 Unknown X 110 Unknown X 113 Unknown X 153 Unknown X 201 Unknown X 215 Unknown X 321 Unknown X

MONTHLY TOTALS 3 11 23 71 132 15 Table 6. Three-year total number of epigeous macrofungal sporocarp_s :r_e~peci~~_presen!_in each plot in_the study area. Species 1:1 1:2 1:3 2:1 2:2 2:3 3:1 3:2 3:3 4:1 4:2 4:3 5:1 5:2 5:3 Boletales Boletus subtomentosus L. Boletus sp. (#313) 2 Chroogomphus corallinus O.K. Miller & Watl. 2 Leccinum aurantiacum (Bull. ex St. Am.) S. F. Gray1 1 Suillus granulatus (L.:Fr.) Roussel 3 4 3 Suillus pictus (Peck) A.H. Sm. & Theirs 5 2 5 1 6 Suillus placidus (Bonord.) Sing. 5 Suillus subalutaceous (A.H. Sm. & Thiers) 2 A.H. Sm. & Thiers Suillus sp. (#K) 46 21 Suillus sp. (#Z) Suillus sp. (#37) 2 Suillus sp. (?) (#Y) 2 4 3 15 Tylopilus felleus (Bull.:Fr.) Karst. 2

Agaricales Amanita ceciliae (Berk. & Br.) Bas Amanita flavoconia Atk. 14 Amanita fulva (Schaeff.) Pers. 16 2 Amanitafulva (?) (Schaeff.) Pers. 9 Amanita porphyria (Alb. & Schw.:Fr.) Mlady Amanita vagina/a (Bull. ex Fr.) Que!. 2 Amanita virosa (Fr.) G. Berto!. 2 1 2 2 Amanita sp. (#R) 1 Amanita sp. (#206) Armillaria mellea complex (Vahl:Fr.) Kumm. 3 33 21 11 3 5 5 89 Chrysomphalina sp. (#14) 98 Clitocybe clavipes (Pers.:Fr.) Kumm. 5 Clitocybe s inopicoides Peck 11 8 Clitocybe vilescens (?) Peck 12 00 \0 Table 6. Continued SEecies 1:1 1:2 1:3 2:1 2:2 2:3 3:1 3:2 3:3 4:1 4:2 4:3 5:1 5:2 5:3 Clitocybe sp. (#213) Collybia butyracea (Bull.:Fr.) Kumm. 2 13 7 8 Collybia confluens (Pers.:Fr.) Kumm. 1 5 273 Collybia dryophila (Bull.:Fr.) Kumm. 37 6 8 20 86 9 3 5 2 34 Collybia luteifolia Gill.3 2 9 1 1 Collybia maculata (Alb. & Schw.:Fr.) Kumm. 6 Collybia proxilla (Fr.) Gill. Coprinellus angulatus Peck 14 14 2 Entoloma majaloides Orton 10 Hygrocybe acutoconica var. microspora (?) 7 Hesler & A.H. Sm. Hygrocybe conica (Scop.:Fr.) Kumm. var. conica Hygrocybe laeta (Pers.:Fr.) Kumm. 1 3 7 17 25 30 11 4 Hygrocybe marginata Peck var. marginata 3 2 7 Hygrocybe miniata (Fr.) Kumm. 19 9 4 20 13 Hygrocybe miniata (Berk. & Br.) E. Arnolds 3 6 3 var. mollis (?) Hygrocybe miniat a (?) (Fr.) Kumm. 4 Hygrophorus angustifolius (Murr.) Hesler & 27 3 A.H.Sm. Hygrophorus camarophyllus (Alb. & Schw.:Fr.) 3 Dumee, Grandjean & Maire Hygrophorus russula (Scop.:Fr.) Quel. 3 Hygrophorus sp. (?) (#195) 2 Hygrophorus sp. (?) (#338) 3 Hypholomafasciculare (Huds.:Fr.) Kumm. 274 Hypholomafasciculare (?) (Huds.:Fr.) Kumm. 14 Laccaria bicolor (Maire) Orton 2 Laccaria laccata (Scop.:Fr.) Berk. & Br. 38 182 30 13 2 1 20 4 13 Leptoniaformosus (Fr.) Quel. Leptonia fulvus (?) (Orton) Mos. 6 \0 Leptonia sericellum (Bull. ex Fr.) Kumm. 2 0 Table 6. Continued seecies 1:1 1:2 1:3 2:1 2:2 2:3 3:1 3:2 3:3 4:1 4:2 4:3 5:1 5:2 5:3 Leptonia sp. (#43) 19 Leptonia sp. (#145) 3 1 3 3 Leptonia sp. (#199) 3 Marasmius androsaceus (L.:Fr.) Fr. many many Marasmius sp. (#111) 8 121 Mycena epipterygia var. pelliculosa (?) (Fr.) Maas G. 4 Mycena griseoviridis A.H. Sm. Mycena griseoviridis (?) A.H. Sm. 2 Mycena sanguinolenta (Alb. & Schw.:Fr.) Kumm. 30 47 52 3 4 2 2 Mycena sp. (#102) 4 1 3 16 7 7 5 14 5 50 Mycena sp. (# 179) 17 Mycena sp. (#307) 3 Mycena sp. (#319) Myxomphalia maura (Fr.) Hora 20 24 82 Myxomphalia maura (?) (Fr.) Hora 6 Nolanea nitens (?) (Vel.) K. & R. 2 Nolanea sp. (#116) Pho/iota highlandensis (Peck) A.H. Sm. & Hesler 7 1 28 217 13 Tricholoma apium (?) Schaeff. Tricholoma columbetta (Fr.) Kumm. 10 3 9 Tricholomaflavovirens (Pers.:Fr.) Lundell Tricholoma imbricatum (?) (Fr.:Fr.) Kumm. Tricholoma magnivelare (Peck) Redhead 1 Tricholoma saponaceum (Fr.:Fr.) Kumm. 3 Tricholoma sejunctum (Sow.:Fr.) Quel. 1 4 12 6 Tricholomopsis rutilans (Schaeff.:Fr.) Sing. 3 Xeromphalina cauticinalis Kiihn. & Maire 4 3 1 2

Cortinariales Cortinarius alboviolaceus (Pers.:Fr.) Fr. 24 2 18 8 9

Cortinarius armillatus (Fr.:Fr.) Fr. 1 55 1 ,___.\0 Table 6. Continued Seecies 1:1 1:2 1:3 2:1 2:2 2:3 3:1 3:2 3:3 4:1 4:2 4:3 5:1 5:2 5:3 Cortinarius bolaris (Pers.:Fr.) Fr. 2 9 10 7 2 Cortinarius cinnamomeus (L.:Fr.) Fr. 1 13 Cortinarius leucopus (?) (Bull. ex Fr.) Fr. 2 1 7 19 2 6 11 42 9 Cortinarius lividoochraceus (?) (Berk.) Berk. 2 Cortinarius multiformis var. coniferarum Mos. 3 Cortinarius obliquus Peck 3 1 1 6 1 11 Cortinarius obtusus (?) (Fr.) Fr. 16 2 3 4 Cortinarius rubricosus (?) Fr. 4 Cortinarius semisanguineus (Fr.) Gill. 3 14 34 Cortinarius sphaerosporus Peck Cortinarius subtorvus (?) Lamoure 4 6 Cortinarius tenebricus (?) Favre 1 4 7 5 Cortinarius trivia/is Lange 2 Cortinarius sp. (# 1/27) 3 1 2 2 8 8 10 Cortinarius sp. (# 15) 2 1 2 Cortinarius sp. (#23b) 2 Cortinarius sp. (# 152) Cortinarius sp. (# 193) Cortinarius sp. (# 196) 9 7 Cortinarius sp. (#317) 7 Cortinarius sp. (#332) 3 Cortinarius sp. (?) (#214) 2 5 4 3 Cortinarius sp. (?) (#318) 1 Hebeloma crustuliniforme (Bull.) Quel. 3 3 3 Hebeloma sinapizans (?) (Paul.) Gill. Hebeloma sp. (?) (#167) 2 Inocybe lacera (Fr.) Kumm. 12 Inocybe leptophylla Atk. 3 5 Inocybe proximella (?) Karst. 7 Inocybe sp. (#24/335) 12 1 14 1 \0 Inocybe sp. (#25) 1 N Table 6. Continued S~ecies 1:1 1:2 1:3 2:1 2:2 2:3 3:1 3:2 3:3 4:1 4:2 4:3 5:1 5:2 5:3 Inocybe sp. (#214) Naucoria sp. (#312) Phaeocollybia christinae (Fr.) Heim Phaeocollybiajennyae (Karst.) Heim 20 Rozites caperatus (Pers.:Fr.) Karst. 6 1 2 3

Russulales Lactarius a/finis var. a/finis Peck 5 2 Lactarius camphoratus (Bull.:Fr.) Fr. 1 47 5 2 2 Lactarius deceptivus Peck 2 7 Lactarius hibbardae var. hibbardae Peck Lactarius lignyotus Fr 2 Lactarius sordidus Peck 2 Lactarius subvellereus var. subvellereus Peck 12 Lactarius torminosus (Schaeff.:Fr.) Pers. Lactarius uvidus (Fr.) Fr. var. uvidus Lactarius vinaceorufescens A.H. Sm. 19 1 1 34 7 43 1 6 22 25 Lactarius sp. (#AA/E/26) 11 6 7 7 1 26 1 4 1 11 1 Russula bicolor (?) Burl. 1 4 Russula brevipies Peck 2 1 1 9 Russula brunneola Burl. 4 Russula cyanoxantha (Schaeff.) Fr. f. cyanoxantha 2 1 2 Russula decolorans (?) (Fr.) Fr. 2 Russula elaeodes (Bres.) Bon Russula faginea (?) Romagn. Russula fragilis (Pers. :Fr.) Fr. 2 6 124 1 5 10 14 Russulafragrantissima Romagn. 3 3 9 3 Russula incarnaticeps (?) Murrill Russula paludosa Britz. 4 Russula paludosa (?) Britz. (# 157) 3 \0 Russula paludosa (?) Britz. (#337) 2 1 2 2 VJ Table 6. Continued Species 1:1 1:2 1:3 2: 1 2:2 2:3 3: 1 3:2 3:3 4: 1 4:2 4:3 5: 1 5:2 5:3 Russula sanguinea (Bull.) Fr. Russula sanguinea (?) (Bull.) Fr. 2 1 1 8 10 3 7 1 5 Russula sp. (#J) Russula sp. (#38) 4 Russula sp. (#171) 3 Russula sp. (#200) 2 Russula sp. (#314) 1 4 4 Russula sp. (#325) Russula sp. (#339) Russula sp. (#340) Russula sp. (#341) 2 Russula sp. (#342)

Unknown species: collection number & year # AIB, 1999 # C (Cortinarius sp. ?), 1999 # P, 1999 # Q, 1999 # S, 1999 39 #U, 1999 8 # V, 1999 1 4 3 # w, 1999 # X, 1999 2 # 16, 1999 1 4 # 17, 1999 2 # 18, 1999 # 28, 1999 3 #32, 1999 3 # 35, 1999 15 # 39, 1999 3 \0 # 42, 1999 ..J:::. Table 6. Continued Species 1: 1 1:2 1:3 2: 1 2:2 2:3 3: 1 3:2 3:3 4: 1 4:2 4:3 5: 1 5:2 5:3 # 110, 2000 6 # 113, 2000 2 # 153, 2000 # 201, 2000 1 4 #215,2000 2 # 321, 2001 2

TOTALSPOROCARPSPERPLOT 233 269 67 159 4 584 169 277 246 170 245 266 358 490 618 TOT AL SPECIES PER PLOT 34 24 12 20 2 46 22 11 29 32 35 50 25 46 30

\0 Vl 96

APPENDIXC.

MAPS 97

:: IAnit 4 c.oV\+rol riots * :: ~"'it 5" contrDI plots

Fig. 1. United States Geological Survey topographic map showing details of the study site in relation to the Boundary Waters Canoe Area Wilderness (BWCA W). 98

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Fig. 2. Location of the three burned study units and the control plots of Units 4 and 5 in relation to the Echo Trail, North and South Hegman lakes, and the Boundary Waters Canoe Area Wilderness (BWCA W). 99

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Fig. 3. Unit 1 detail. (Note the exclusion area where one of the plots of Unit 5 was located.) 100

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Fig. 4. Unit 2 detail. The gravel pit is at the northern boundary of Unit 2. 101

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Fig. 5. Unit 3 detail. Unit 3 was accessed by the old skid road that angled southwest from the Echo Trail and by the improved fire line that skirted the northeast edge of the pond.